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ISSUEISSUE 42,42, PUBLISHEDPUBLISHED 2525 APRILAPRIL 20202020 AustralasianAustralasian JournalJournal ofof HerpetologyHerpetology

ISSN 1836-5698 (Print) ISSN 1836-5779 (Online) CONTENTS ON PAGE 2 2 Australasian Journal of Herpetology

Australasian Journal of Herpetology Issue 42, 25 April 2020. Contents

From a putative new taxon to a mutt! Formal descriptions of three new genetically divergent Mountain Pygmy Possums from Victoria and New South Wales closely associated with Burramys parvus Broom, 1896. ... Raymond T. Hoser, 3-10. Small and overlooked … six new species of Pygmy Possum, Genus Cercartetus Gloger, 1841 sensu lato from the Australasian bioregion. ... Raymond T. Hoser, 11-22. A new subgenus, three new species and one new subspecies of Ring-tailed Possums (Marsupialia: Petauridae) from the north of Australia. ... Raymond T. Hoser, 23-30. Small, easily overlooked and in decline. Potoroos in Eastern Australia. A formal division of the genus Potorous Desmarest, 1804 (Marsupialia Potoroidae) and the description of a new species from south-east Queensland. ... Raymond T. Hoser, 31-37. A new subspecies of Yellow-bellied Glider (Marsupialia: Petauridae) from far north Queensland, Australia. ... Raymond T. Hoser, 38-41. The ability to conserve a threatened species begins when they are named! New species of Rock Wallaby (Marsupialia: Macropodidae: Petrogale) from northern Australia. ... Raymond T. Hoser, 42-49. An overdue refinement of the taxonomy of the Australian Ring Tailed Dragons, Genus Ctenophorus Fitzinger, 1843, Subgenus Tachyon Wells and Wellington, 1985, including the formal descriptions of eight new species. ... Raymond T. Hoser, 50-64.

Front cover photo: Raymond Hoser. Alpine habitat for Mountain Pygmy Possums (Burramys sp.) adjacent to Mount Hotham, Victoria.

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All Intellectual Property (IP) rights are reserved, including in relation to all IP generated by the journal in Hoser 2016 - Australasian Journal of Herpetology 40. terms of both authors, publisher and the like. Online journals (this issue) do Availablenot appear for online a month at after www.herp.net the actual and listed publication date of the printed journals. MinimumCopyright- print run Kotabi of first Publishingprintings is always - All at rightsleast fifty reserved hard copies. Australasian Journal of Herpetology 3 Australasian Journal of Herpetology 42:3-10. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

LSID urn:lsid:zoobank.org:pub:B5F27303-4481-4E44-9EEF-26747C600803

RAYMOND T. HOSER LSID urn:lsid:zoobank.org:author:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE 488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 28 March 2020, Accepted 2 April 2020, Published 25 April 2020.

ABSTRACT The iconic Mountain Pygmy Possum Burramys parvus Broom, 1896, was originally described from Pleistocene fossil material obtained near Taralga, New South Wales. Since then, living specimens of the same putative species have been found in alpine areas of New South Wales and Victoria broadly coinciding with the major ski resorts in each state. Presumption that these living animals are conspecific with the type material in the Australian Museum in Sydney has been made by all publishing zoologists to date in the absence of genetic evidence. Molecular evidence published by Osborne et al. (2000), implied divergences in the three main extant populations of up to 680,000 years before present, which they stated “conflicts with conclusions reached by Ride and Davies (1997), that the extant populations of B. parvus are remnants of a species that had a more contiguous distribution as recently as 18 000 years ago.” Significantly Mitrovski et al. (2007) wrote “From a conservation perspective, all three areas where B. parvus are found should be considered as separate gene pools” implying the need for each to have taxonomic recognition. Strahan (1988) also suggested subspecies level recognition for the regional populations of putative B. parvus. Following on from these works, this paper takes a conservative position and formally names all three known extant races as subspecies, separate from the fossil type material. Unfortunately, in 2008, the government-owned “Zoos Victoria” business cross-bred Mount Hotham and Mount Buller animals at their facility at Healesville and along with translocation of specimens from Mount Hotham to Mount Buller had by 2012 successfully created a cohort of at least 50% of trapped wild Mount Buller specimens (9 of 18) as hybrids (Menkhorst et al. 2016), signifying that as of 2020, the pure Mount Buller lineage is probably extinct. In other words all Burramys at Mount Buller are effectively Mutts! Formal naming of the pure Mount Buller form (based on previously lodged material in the Museum of Victoria) and the Bogong Plains form (including Mount Hotham) and the main population from the Alps in New South Wales will hopefully help to prevent any other ill-conceived attempts to further mix these taxonomically distinct and allopatric populations. Keywords: Mammals; Marsupial; taxonomy; nomenclature; Pygmy Possum; Burramys; parvus; Mount Buller; Mount Hotham; Victoria; New South Wales; new subspecies; hosersbogensis; timdalei; scottyjamesi.

INTRODUCTION as subspecies. The iconic Mountain Pygmy Possum Burramys parvus Broom, I go further and also formally name the northern form from the 1896, was originally described from Pleistocene fossil material Kosciuszko region of New South Wales as another subspecies obtained at Wombeyan Caves near Mittagong in the New South on the basis of morphological differences in the jaw between all Wales Southern Highlands in 1895 (Broom 1896). living forms and the type fossil material (see also Broom 1896 A living animal, believed to be of the same species was who used the type material for examination, and Brammall and discovered at Mount Hotham Ski Resort in Victoria in 1966, and Archer 1997, who used a Victorian specimen for review populations of the same putative species were subsequently purposes). discovered in the Kosciuszko region of NSW in 1970 and at the Unfortunately, in 2008, and following an internal review of the Mount Buller Ski resort in 1996 (Calaby et al. 1971, Mansergh Mount Buller population, the government-owned “Zoos Victoria” and Broome 1994, Heinz and Williams 1998). business in conjunction with the State Wildlife Department that controls them, caught a number of wild specimens and cross- Further fossil remains attributed to the same putative species bred Mount Hotham and Mount Buller animals at their facility at were recovered from caves at Buchan Victoria and south-west of Healesville. Sydney in New South Wales (Jenolan Caves) and were treated as suggesting that at the height of the last Pleistocene glacial Along with translocation of specimens from Mount Hotham to period (ca. 20,000 years bp) a single species of Mountain Mount Buller, they had by 2012 successfully created a cohort of Pygmy-possum had a much wider distribution around the at least 50% of trapped wild Mount Buller specimens (9 of 18) as snowline of south-eastern Australia (Ride and Davies 1997). hybrids (Menkhorst et al. 2016), signifying that as of 2020, the It has been generally assumed that since that glacial maximum pure Mount Buller lineage is probably extinct. receded, the range of the putative species has been contracting In other words all Burramys at Mount Buller are effectively Mutts! with a gradually warming climate and receding snowline The alleged basis of the action by “Zoos Victoria” for creating the (Caughley 1986; Mansergh and Broome 1994). Mutt population at Mount Buller was due to a potentially In line with the preceding, the presumption that these living misguided belief that the Mount Buller population was in animals are conspecific with the type material in the Australian immediate danger of extinction and in their view too small to be Museum in Sydney has been made by all publishing zoologists viable in the long term and therefore needed greater genetic to date in the absence of genetic evidence. diversity in the form of an infusion from the Mount Hotham However molecular evidence published by Osborne et al. population. (2000), implied divergences in the three main extant populations The scientific basis for the belief that the Mount Buller of up to 680,000 years before present, which they stated population would otherwise rapidly and inevitably die out was “conflicts with conclusions reached by Ride and Davies (1997), tenuous at best and even if it had a sound basis of scientific fact that the extant populations of B. parvus are remnants of a (which it did not), a better situation would have been to attempt, species that had a more contiguous distribution as recently as using the vast resources available to the government, to save 18 000 years ago.” that population without resorting to hybridising with the These findings also effectively ruled out the idea that the three genetically divergent and allopatric Mount Hotham population. major populations were contiguous at the time of the most Associated with this would be the risk of failure, after which, if recent glacial maxima, this idea in effect being based solely on the Mount Buller population actually did die out, the government crude, temperature-based climate models. people could then consider what to do in the way of successful Identifying the physical barriers to movement for the populations long term re-reintroductions from elsewhere of the same or of the putative species is not easy, as 20,000 years before similar species, viz the Mount Hotham animals. present, not only would vegetation regimes have been different, The worst case scenario would be the same projected end-point, but so too would have been species compositions in intervening viz over-running the place with Mount Hotham animals, but this areas, including competing and predatory ones. second best outcome to preservation of the unique lineage, may Noting that extant putative B. parvus have strict habitat have been wholly avoided. requirements beyond those of maximum temperature tolerance, I make it clear that the better case scenario would have been I did a simple inspection of topographical data to see if in fact long term survival of the Mount Buller population as a genetically putative B. parvus from the Kosciuszko region of NSW would in pure gene pool that to date had survived many tens of fact have a reasonable prospect of surviving in and passing thousands of years in isolation from the others and if reasonably through the areas between known current locations and the well managed, would have continued to do so indefinitely. locations of Pleistocene deposits south-west of Sydney, NSW. Formal naming of the pure Mount Buller form (based on types in The need for rocky boulder habitat of cold temperatures, either the Museum of Victoria) and the Bogong Plains form (including due to habitat specific requirements or predator avoidance Mount Hotham) will hopefully prevent introduction of other strategies, rendered most of the intervening region unsuitable, subspecies into the wholly viable, allopatric and distinct even in times of glacial maxima, furthering my belief that type B. population of the Victorian Bogong Plains and draw attention to parvus is almost certainly not even be conspecific with the living the fact that preservation of the species in their natural form animals being treated as the same species. should be the first option in wildlife conservation and not the creation of Mutts! In any event, even if the northern (New South Wales) population were to be conspecific with type B. parvus, it is clear based on The naming of the Victorian populations in line with the above is published genetic analysis (e.g. Osborne et al. 2000), that both due to the obvious contention that they are most geographically the main Victorian populations are sufficiently divergent from divergent from the New South Wales stock and the fossil type these and from one another, to be regarded as subspecies on from north of there (presumed for the moment to be conspecific the basis they are morphologically divergent, allopatric, at the subspecies level) and so by simple logic, were at the genetically divergent for thousands of years and have no gene outset of the investigation being the divergent specimens in flow between each other. need of formal taxonomic recognition. Significantly Mitrovski et al. (2007) wrote “From a conservation MATERIALS, METHODS AND RESULTS perspective, all three areas where B. parvus are found should be Before a decision is made to name any new taxon, reasonable considered as separate gene pools” further implying the need for steps must be taken to ensure that it is justified on all relevant each to have taxonomic recognition. grounds, including that it is morphologically, genetically and This paper does exactly that and formally names the two reproductively isolated from their nearest relative. Once these southern races, being most distant from the type fossil material have been ascertained, the decision as to at what nomenclatural

level needs to be assigned to ensure stability of names and was in fact of the same species and genetic lineage. utility in the Zoological community. The connecting line also includes significant rocky areas that That is, should the reproductively isolated and morphologically would clearly be of suitable habitat in times of glacial maxima, divergent entities be labelled as subspecies, full species, or further supporting the above contention. potentially higher level again. There is no extant high altitude bioregion at Buchan that in any Key literature relevant to the taxonomic and nomenclatural way has endemism not seen at Mount Hotham, Buller or conclusions within this paper include Kosciuszko, further implying that the Buchan specimen is in fact Anonymous (2005), Bannister et al. (1988), Broom (1895, an outlier from a colder geological period as opposed to being a 1896a, 1896b). Broome (2001a, 2001b), Broome and Geiser taxonomically discrete entity. (1995), Broome et al. (2012), Calaby et al. (1971), Caughley The molecular phylogeny of Pepper et al. (2018) confirmed that (1986), Dimpel and Calaby (1972), Fleming (1985), Geiser and putative Eulamprus kosciuskoi (Kinghorn, 1923) from the Blue Broome (1991, 1993), Gray (1841, 1845), Green (2003, 2008), Mountains region of New South Wales, was in fact a different Gullan and Norris (1984), Heinze and Olejniczak (2000), Heinze species-level taxon to the nominate form from a 5,000 feet and Williams (2008), Heinze et al. (2004), Hoser (1991, 2019a, elevation on Mount Kosciuszko. 2019b), Kerle (1984), Körtner and Geiser (1998), Mansergh and The Blue Mountains species was in fact formally identified and Broome (1994), Mansergh and Scotts (1989, 1990), Mansergh named by Wells and Wellington (1984) as Eulamprus and Walsh (1983), Mansergh et al. (1989, 1990, 2010), leuraensis. Menkhorst and Knight (2004), Menkhorst et al. (2016), Mitrovski See also the molecular phylogeny for the Mountain Dragon et al. (2005, 2007, 2008), Ng et al. (2013), Osborne and species complex, all treated by most authors as a single putative Christidis (2002), Osborne et al. (2000), Pepper et al. (2018), species, being Rankinia diemensis (Gray, 1841), found by Ng et Ride (1956), Ride et al. (1999), Rosengren and Peterson (1989), al. (2013) to be a complex of six divergent lineages. Sanecki et al. (2006), Schulz et al. (2012a, 2012b), Shi (2012), Hoser (2015) determined that the antiquity of divergence, Smith and Broome (1992), Van der Ree et al. (2009), Weeks et morphological differences and geographical isolation and al. (2011, 2012), Wells and Wellington (1984, 1985) and sources allopatry necessitated the recognition of each form as a new cited therein. species and formally named the four unnamed forms. While there is an element of doubt over the conspecfic status of Included in this complex, were one form from Tasmania, three the living putative Burramys parvus and the fossil specimens from Victoria and two from New South Wales, those being one from further north, until this is resolved, it makes sense for the from the Blue Mountains bioregion and another from an isolated southern specimens to be treated as subspecies rather than pocket further west. separate species-level entities. In terms of these clades of Mountain Dragons formally named In the event that they are ultimately regarded as different by Hoser (2015), Ng et al. (2013) had written: “there were deep species, one or more of the subspecies formally named herein genetic divergences between Victorian and NSW samples ([8.2 can be simply elevated to full species, without significantly % mtDNA uncorrected sequence divergence), probably dating to destabilizing the existing in use nomenclature. the late Miocene-Pliocene. All relevant previously published material was examined, Multiple phylogeographic studies have also found deep genetic including that cited herein and sources within those publications. breaks in southern NSW (e.g., Chapple et al. 2005; Symula et Live and dead specimens as well as available bone specimens, al. 2008).” were examined as was other relevant material, including past climate data for Victoria and New South Wales, past vegetation The dry habitat barriers referred to by Chapple et al. (2005), maps, previously published literature on these matters, including would of course have similarly prevented Burramys populations phylogenic treatments of other species groups distributed or in south-east New South Wales from mixing as well, even in range restricted in high altitude areas of south-east Australia times of cooler and drier climates. and so on in order to gauge likely spread of extant populations in In simple terms, even in times of glacial maxima, there would the recent geological past. not have been an obvious conduit for Snowy Mountains (NSW) Genetic data published was also scrutinized as were papers and Blue Mountains (NSW) populations of Burramys to have dealing with the biology, captive breeding and other relevant mixed. facts about the relevant species. On the basis of the preceding one can only expect that all three In summary, as inferred already, the genetic evidence was main populations of living putative Burramys parvus are quite definitive in that it showed 1/ That the three main living likely a new and different species to the original type material populations of B. parvus had remained distinct and separate from south-west of Sydney, New South Wales! well before the most recent glacial maxima about 20,000 years However in the absence of genetic data from the fossil material, before present and 2/ There is no evidence available, including I have taken the conservative view and named the two southern via the fossil record that the main region between living forms and the NSW form as subspecies of the type fossil specimens and the fossils south-west of Sydney in New South material. Wales, was inhabited by any Burramys in the recent past. In the event that fossil Burramys parvus is later shown to be in I also note that the fossil remains of putative B. parvus of fact the same species as the living putative B. parvus, noting Pleistocene age from Buchan, Victoria are in a very different that there are documented differences in the available skull position to those from near Wombeyan and Jenolan Caves in material from each as seen in the published literature (refer to NSW. Broom 1896 and comparative material from Victoria in Brammall Those two NSW sites are widely divergent from extant and Archer 1997), then the taxonomy and nomenclature in this populations and sit on the edge of the high altitude Blue paper need never change. Mountains escarpment, a significant alpine bioregion in its own If however the fossil B. parvus is subsequently shown to be of a right and with a high degree of alpine endemism (see for different species, then one of the three forms named herein (the example Pepper et al. 2018). first) can be elevated to full species without significantly By contrast, Buchan, Victoria sits on the immediate southern destabilising the taxonomy of all relevant populations. edge of the Victorian Great Dividing Range and is directly Rather than run the risk of species or subspecies becoming connected to the extant New South Wales population in the threatened or extinct due to non-recognition of them, as shown Kosciuszko region of NSW via the Davies Plain Ridge in in Hoser (2019a, 2019b), this paper has been published, even Victoria, or alternatively via the ridgeline of high mountain peaks though it is clearly too late to save the Mount Buller taxon of immediately east of there, inferring that this specimen at least Burramys in its pure form. Hoser 2020 - Australasian Journal of Herpetology 42:3-10. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 6 Australasian Journal of Herpetology

INFORMATION RELEVANT TO THE FORMAL DESCRIPTIONS Putative wild B. parvus will be faced with ever shrinking potential THAT FOLLOW habitat to live in and how larger human populations both in There is no conflict of interest in terms of this paper or the Australia and globally can reduce the likelihood of this conclusions arrived at herein. catastrophe being averted, other than by stopping population Several people including anonymous peer reviewers who increases is unknown. revised the manuscript prior to publication are also thanked as a BURRAMYS PARVUS HOSERSBOGENSIS SUBSP. NOV. relevant staff at museums who made specimens and records LSID urn:lsid:zoobank.org:act:7DE00FBD-90BD-40FA-9E3C- available in line with international obligations. 9AD6E7C55292 In terms of the following formal descriptions, spellings should Holotype: A preserved specimen at the Australian Museum in not be altered in any way for any purpose unless expressly and Sydney, New South Wales, Australia, specimen number exclusively called for by the rules governing Zoological M.36919 collected at the double toilet at Mount Perisher Ski Nomenclature as administered by the International Commission Resort, New South Wales, Australia, Latitude -36.40 S., of Zoological Nomenclature. Longitude 148.38 E. This government-owned facility allows access to its holdings. In the unlikely event two or more newly named taxa are deemed conspecific by a first reviser, then the name to be used and Paratypes: Four preserved specimens at the Australian retained is that which first appears in this paper by way of page Museum in Sydney, New South Wales, Australia, specimen priority and as listed in the abstract keywords. numbers M24919, M.24920, M24921 and M.24922 collected Some material in descriptions for taxa may be repeated for other from Mount Blue Cow, Kosciusko National Park, New South taxa in this paper and this is necessary to ensure each fully Wales, Australia, Latitude -36.32 S., Longitude 148.42 E. complies with the provisions of the International Code of Diagnosis: Living Burramys parvus Broom, 1896 of all three Zoological Nomenclature (Fourth edition) (Ride et al. 1999) as subspecies named herein are formally separated from the fossil amended online since. type material herein treated as a fourth and nominate Material downloaded from the internet and cited anywhere in this subspecies, namely B. parvus parvus. paper was downloaded and checked most recently as of 10 All three newly named subspecies of B. parvus are readily January 2020, unless otherwise stated and was accurate in separated from the fossil type material from near Taralga, New terms of the context cited herein as of that date. South Wales by the fact that the premolars are in proportion, versus slightly oversized in fossil B. parvus type material, which Unless otherwise stated explicitly, colour and other descriptions is evidence and grounds to form the basis that the living forms apply to living adult specimens of generally good health and not are at least different subspecies (also refer to Broom 1896 and under any form of stress by means such as excessive cool, comparative material from Victoria in Brammall and Archer heat, dehydration or abnormal skin reaction to chemical or other 1997). input. Colour descriptions of species refer to fur colour and not skin. Besides being easily separated from one another by genetics as detailed by Osborne et al. (2000), the three newly named living While numerous texts and references were consulted prior to subspecies of B. parvus are in turn readily separated from one publication of this paper, the criteria used to separate the another by colouration as follows: relevant species or subspecies has already been spelt out and/ B. parvus hosersbogensis subsp. nov. from the Kosciusko or is done so within each formal description and does not rely on National Park, New South Wales, Australia are grey-brown material within publications not explicitly cited herein. above, sometimes darker in the mid-dorsal area, continuing to In terms of conservation of each population of each subspecies the top of the head; dark ring around the eye; pale grey-brown or as described below, the comments in Hoser (1991 and 2019a, pale brown below; light brown on the cheeks. There is some 2019b) apply. I note that based on the information provided by white between the eyes and the tip of the snout. Menkhorst (2016) the subspecies Burramys parvus timdalei B. parvus timdalei subsp. nov. from Mount Buller in Victoria is subsp. nov. may have been literally bred out of existence, at yellow-brown above, sometimes darker in the mid-dorsal area, least in its pure form. continuing to the top of the head; dark ring around the eye; light I also note that the document “National Recovery Plan for the brown below and yellow cheeks. Mountain Pygmy-possum Burramys parvus”, (Menkhorst 2016), wholly fails to address the primary root cause of likely declines There is no white between the eyes and the tip of the snout. and extinctions in populations of putative Burramys parvus. Instead this area is a deep yellow. This is the relentless ongoing rapid growth in the human B. parvus scottyjamesi subsp. nov. from the Bogong High Plains population in Australia and environmental pressures this brings. including the Mount Hotham Ski resort is grey-brown above, Of course as a Victorian State Government Employee, sometimes darker in the mid-dorsal area, continuing to the top Menkhorst is not allowed to point out the serious defects in his of the head; dark ring around the eye; cream or white coloured employer’s “Big Australia Policy” which in its most recent form, below and light cream or white on the cheeks. was a population target of 100-125 million people in Australia There is some white between the eyes and the tip of the snout. and one quarter that number for Victoria, as soon as practicable B. parvus hosersbogensis subsp. nov. in life can be seen online (up from the 25 million nationally and 6 million in Victoria 2020). at: On these numbers alone, even if humans were forcibly https://www.flickr.com/photos/alexismarie_meyer/4464197606/ barricaded from the regions where the Burramys parvus live and (being the ski resorts), the marsupials will no doubt die off from https://www.flickr.com/photos/152410663@N05/35729353956/ ingesting ever increasing quantities of pesticides that blow in on their insect food that emanates from farming regions nearby (both last downloaded on 10 January 2020). (Green 2008), that must as a matter of survival for humans, be B. parvus timdalei subsp. nov. in life can be seen online at: farmed with ever increasing intensity. http://www.mtbuller.com.au/uploads/file/MPP_Fact_Sheet.pdf Hence the long-term prognosis for all extant populations of and putative Barramys parvus in the wild is in fact tenuous at best. https://cdn.mtbullercdn.com.au/assets/environment/ I need not mention the likelihood of global warming caused mt_buller_mountain_pygmy-possum_recovery_plan_2011- primarily by ongoing human overpopulation and the threat this 2016_final.pdf brings to the cold climate species caught in islands of habitat in (both last downloaded on 10 January 2020). the form of mountain peaks and uplands. B. parvus scottyjamesi subsp. nov. in life can be seen in Strahan

(1988) on page 168 (both images), on the front cover of Hoser South Wales by the fact that the premolars are in proportion, (1991) as well as on page 217 of that book (different image, and versus slightly oversized in fossil B. parvus type material, which clearly showing the diagnostic features of this subspecies) and is evidence and grounds to form the basis that the living forms images of the same animal are seen at: are at least different subspecies (also refer to Broom 1896 and http://www.edgeofexistence.org/species/mountain-pygmy- comparative material from Victoria in Brammall and Archer possum/ 1997). (last downloaded on 10 January 2020). Besides being easily separated from one another by genetics as As of 2019, according to most publishing authors, there are two detailed by Osborne et al. (2000), the three newly named living extant genera of pygmy possums: Burramys Broom, 1896 and subspecies of B. parvus are in turn readily separated from one Cercartetus Gloger, 1841. Burramys contains only one extant another by colouration as follows: species, the Mountain Pygmy-possum, Burramys parvus, which B. parvus hosersbogensis subsp. nov. from the Kosciusko has a lightly furred tail. National Park, New South Wales, Australia are grey-brown above, sometimes darker in the mid-dorsal area, continuing to The morphologically similar Cercartetus as recognized in 2019 the top of the head; dark ring around the eye; pale grey-brown or consists of four currently recognized species being C. caudatus pale brown below; light brown on the cheeks. There is some (Milne-Edwards, 1877) (in fact at least 6 species), C. Lepidus white between the eyes and the tip of the snout. Thomas, 1888 (in fact two species), C. concinnus (Gould, 1845) B. parvus timdalei subsp. nov. from Mount Buller in Victoria is and C. nanus (Desmarest, 1818). yellow-brown above, sometimes darker in the mid-dorsal area, For evidence and reasons explaining why there are in fact more continuing to the top of the head; dark ring around the eye; light extant species than those listed as recognized above, see brown below and yellow cheeks. Osborne and Christidis (2002) and then see Hoser (2020). The genus Eudromicia Mjöberg, 1916 is resurrected by Hoser There is no white between the eyes and the tip of the snout. (2020) for the species associated with C. caudatus (Milne- Instead this area is a deep yellow. Edwards, 1877) based on a divergence of 16-27 MYA (Osborne B. parvus scottyjamesi subsp. nov. from the Bogong High Plains and Christidis, 2002). including the Mount Hotham Ski resort is grey-brown above, Cercartetus sensu lato species are separated from Burramys by sometimes darker in the mid-dorsal area, continuing to the top having tails that have a feather-like appearance. of the head; dark ring around the eye; cream or white coloured below and light cream or white on the cheeks. Further relevant diagnostic and other information about B. There is some white between the eyes and the tip of the snout. parvus sensu lato can be found on pages 216 and 217 of Hoser (1991). B. parvus hosersbogensis subsp. nov. in life can be seen online Distribution: B. parvus hosersbogensis subsp. nov. is known at: only from the Kosciusko National Park, New South Wales, https://www.flickr.com/photos/alexismarie_meyer/4464197606/ Australia. and Etymology: The name is a take on that of the type locality. This https://www.flickr.com/photos/152410663@N05/35729353956/ is a public toilet at the Perisher Ski Resort, where I relieved (both last downloaded on 10 January 2020). myself with a massive faeces (Australian word is “bog”), making B. parvus timdalei subsp. nov. in life can be seen online at: it the site of “Hoser’s bog”. It may have been some kind of record breaker, but adding the words “potentially record http://www.mtbuller.com.au/uploads/file/MPP_Fact_Sheet.pdf breaking” to the scientific name would have made it excessively and long as would have been “Hoser’s massive bog”! Hence the https://cdn.mtbullercdn.com.au/assets/environment/ name “hosersbogensis”. mt_buller_mountain_pygmy-possum_recovery_plan_2011- There is also a swamp or “bog nearby. 2016_final.pdf (both last downloaded on 10 January 2020). In any event, this name will be easily remembered by Australian zoologists and lay people alike and draw people’s attention to a B. parvus scottyjamesi subsp. nov. in life can be seen in Strahan threatened taxon. (1988) on page 168 (both images), Thankfully I did not use this toilet curing the Coronvirus epidemic on front cover of Hoser (1991) as well as on page 217 of that of 2020, because for some months in early 2020, panic buying book (different image, and clearly showing the diagnostic by people stripped shop shelves bare of toilet paper and most of features of this subspecies) and images of the same animal at the population in Australia was physically unable to obtain the http://www.edgeofexistence.org/species/mountain-pygmy- commodity. possum/ BURRAMYS PARVUS TIMDALEI SUBSP. NOV. (last downloaded on 10 January 2020). LSIDurn:lsid:zoobank.org:act:7805FD69-CDEB-49BA-A472- As of 2019, according to most publishing authors, there are two C9225440B312 extant genera of pygmy possums: Burramys Broom, 1896 and Holotype: A preserved specimen at the National Museum of Cercartetus Gloger, 1841. Burramys contains only one extant Victoria, Melbourne, Victoria, Australia, specimen number species, the Mountain Pygmy-possum, Burramys parvus, which C37334, collected at Mount Buller Ski Resort, Victoria, Australia, has a lightly furred tail. Latitude -37.13 S., Longitude 164.45 E. This government-owned The morphologically similar Cercartetus as recognized in 2019 facility allows access to its holdings. consists of four currently recognized species being C. caudatus Paratypes: Two preserved specimens at the National Museum (Milne-Edwards, 1877) (in fact at least 6 species), C. Lepidus of Victoria, Melbourne, Victoria, Australia, specimen number Thomas, 1888 (in fact two species), C. concinnus (Gould, 1845) C37335 and C37336, collected at Mount Buller Ski Resort, and C. nanus (Desmarest, 1818). Victoria, Australia, Latitude -37.13 S., Longitude 164.45 E. For evidence and reasons explaining why there are in fact more extant species than those listed as recognized above, see Diagnosis: Living Burramys parvus Broom, 1896 of all three Osborne and Christidis (2002) and then see Hoser (2020). subspecies named herein are formally separated from the fossil type material herein treated as a fourth and nominate The genus Eudromicia Mjöberg, 1916 is resurrected by Hoser subspecies, namely B. parvus parvus. (2020) for the species associated with C. caudatus (Milne- Edwards, 1877) based on a divergence of 16-27 MYA (Osborne All three newly named subspecies of B. parvus are readily and Christidis, 2002). separated from the fossil type material from near Taralga, New Cercartetus sensu lato species are separated from Burramys by Hoser 2020 - Australasian Journal of Herpetology 42:3-10. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 8 Australasian Journal of Herpetology having tails that have a feather-like appearance. below and light cream or white on the cheeks. Cercartetus species are separated from Burramys by having There is some white between the eyes and the tip of the snout. tails that have a feather-like appearance. B. parvus hosersbogensis subsp. nov. in life can be seen online Further relevant diagnostic and other information about B. at: parvus sensu lato can be found on pages 216 and 217 of Hoser https://www.flickr.com/photos/alexismarie_meyer/4464197606/ (1991). and Distribution: B. parvus timdalei subsp. nov. are known only https://www.flickr.com/photos/152410663@N05/35729353956/ from the immediate environs of the Mount Buller Ski Resort, Victoria, Australia, with main populations centred on the (both last downloaded on 10 January 2020). Federation and Fanny’s Finnish ski run areas and associated B. parvus timdalei subsp. nov. in life can be seen online at: boulder fields. http://www.mtbuller.com.au/uploads/file/MPP_Fact_Sheet.pdf Unfortunately the surviving population has been infected with and specimens of the Mount Hotham lineage (Menkhorst et al. https://cdn.mtbullercdn.com.au/assets/environment/ 2016), meaning that pure B. parvus timdalei subsp. nov. are as mt_buller_mountain_pygmy-possum_recovery_plan_2011- of 2020 almost certainly extinct. 2016_final.pdf Etymology: Named in recognition of Tim Dale of Warrandyte, (both last downloaded on 10 January 2020). Victoria, Australia who is a local snowboarding legend at Mount B. parvus scottyjamesi subsp. nov. in life can be seen in Strahan Buller, Victoria and has been for some years in recognition for (1988) on page 168 (both images), his logistical services aiding wildlife conservation by providing on front cover of Hoser (1991) as well as on page 217 of that relevant services to Snakebusters: Australia’s best reptile book (different image, and clearly showing the diagnostic shows, being the only hands on reptile shows in Australia that features of this subspecies) and images of the same animal at educate by letting people hold the animals. http://www.edgeofexistence.org/species/mountain-pygmy- BURRAMYS PARVUS SCOTTYJAMESI SUBSP. NOV. possum/ LSID urn:lsid:zoobank.org:act:269AA0C7-FE7D-4445-9696- (last downloaded on 10 January 2020). 70DAE60B8DDE As of 2019, according to most publishing authors, there are two Holotype: A preserved specimen at the National Museum of extant genera of pygmy possums: Burramys Broom, 1896 and Victoria, Melbourne, Victoria, Australia, specimen number Cercartetus Gloger, 1841. Burramys contains only one extant C27309 collected from the eastern slope of Mount Higginbotham species, the Mountain Pygmy-possum, Burramys parvus, which at the Mount Hotham Ski Resort, Victoria, Australia, Latitude - has a lightly furred tail. 36.98 S., Longitude 147.15 E. The morphologically similar Cercartetus as recognized in 2019 This government-owned facility allows access to its holdings. consists of four currently recognized species being C. caudatus Paratype: A preserved specimen at the National Museum of (Milne-Edwards, 1877) (in fact at least 6 species), C. Lepidus Victoria, Melbourne, Victoria, Australia, specimen number Thomas, 1888 (in fact two species), C. concinnus (Gould, 1845) C26917 collected from the western slope of Mount and C. nanus (Desmarest, 1818). Higginbotham at the Mount Hotham Ski Resort, Victoria, For evidence and reasons explaining why there are in fact more Australia, Latitude -36.98 S., Longitude 147.15 E. extant species than those listed as recognized above, see Diagnosis: Living Burramys parvus Broom, 1896 of all three Osborne and Christidis (2002) and then see Hoser (2020). subspecies named herein are formally separated from the fossil The genus Eudromicia Mjöberg, 1916 is resurrected by Hoser type material herein treated as a fourth and nominate (2020) for the species associated with C. caudatus (Milne- subspecies, namely B. parvus parvus. Edwards, 1877) based on a divergence of 16-27 MYA (Osborne All three newly named subspecies of B. parvus are readily and Christidis, 2002). separated from the fossil type material from near Taralga, New Cercartetus sensu lato species are separated from Burramys by South Wales by the fact that the premolars are in proportion, having tails that have a feather-like appearance. versus slightly oversized in fossil B. parvus type material, which Cercartetus species are separated from Burramys by having is evidence and grounds to form the basis that the living forms tails that have a feather-like appearance. are at least different subspecies (also refer to Broom 1896 and Further relevant diagnostic and other information about B. comparative material from Victoria in Brammall and Archer parvus sensu lato can be found on pages 216 and 217 of Hoser 1997). (1991). Besides being easily separated from one another by genetics as Distribution: B. parvus scottyjamesi subsp. nov. is restricted to detailed by Osborne et al. (2000), the three newly named living the Bogong High Plains and associated mountains in north-east subspecies of B. parvus are in turn readily separated from one Victoria, Australia. another by colouration as follows: Etymology: Named in honour of Scotty James of Warrandyte, B. parvus hosersbogensis subsp. nov. from the Kosciusko Victoria, Australia, in recognition for his services for National Park, New South Wales, Australia are grey-brown snowboarding worldwide. He was the flag bearer for Australia at above, sometimes darker in the mid-dorsal area, continuing to the 2018 Winter Olympics, where he won a bronze medal in the top of the head; dark ring around the eye; pale grey-brown or halfpipe. He has won numerous titles since and has inspired pale brown below; light brown on the cheeks. There is some countless young people to get out of their homes and to enjoy white between the eyes and the tip of the snout. the outdoor environment in sport, which in turn encourages B. parvus timdalei subsp. nov. from Mount Buller in Victoria is people to want to do what is needed to preserve and enhance yellow-brown above, sometimes darker in the mid-dorsal area, the world’s natural assets (except when locked in their homes continuing to the top of the head; dark ring around the eye; light with Coronavirus inspired government “lock downs” like in 2020). brown below and yellow cheeks. REFERENCES CITED There is no white between the eyes and the tip of the snout. Anon. (2005). Recovery Plan for the Mountain Pygmy-possum Instead this area is a deep yellow. Burramys parvus on Mt Buller, Victoria. Mt. 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and Walton, D. M. 1988. Zoological Catalogue of Australia, Green, K. 2008. Migratory Bogong Moths (Agrotis infusa) Volume 5, Mammals. Australian Government Publishing Service, transport arsenic and concentrate it to lethal effect by aestivating Canberra, ACT, Australia:X+274 pp. gregariously in alpine regions of the Snowy Mountains of Brammell, J. and Archer, M. 1997. A new Oligocene-Miocene Australia. Arctic, Antarctic, and Alpine Research 40:74-80. species of Burramys (Marsupialia, Burramyidae) from Gullan, P. and Norris, K. 1984. The habitat of Burramys parvus Riversleigh, northwestern Queensland. Memoirs of the (Broom) in Victoria. pp 417-421 in Smith, A. P. and Hume, I. D. Queensland Museum 41(2):247-268. (eds), Possums and Gliders. Surrey Beatty and Sons, Chipping Broom, R. 1895. On a new fossil mammal allied to Norton, NSW, Australia. Hypsiprymnus but resembling in some points the Heinze, D. A. and Olejniczak, A. M. 2000. First observations of Plagiaulacidae. Abstr. 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M. conservation and ski resorts in Kosciuszko National Park, (eds.), Surrey Beatty and Sons, Chipping Norton, Australia. Australia. Biological Conservation 102:309-323. Hoser, R. T. 1991. Endangered Animals of Australia. Pierson Broome, L. S. 2001b. Seasonal activity, home range and habitat Publishing, Mosman, NSW, Australia:240 pp. use of the mountain pygmy-possum Burramys parvus Hoser, R. T. 2015. Australian agamids: Eighteen new species (Marsupialia: Burramyidae) at Mt Blue Cow, Kosciuszko National from the genera Amphibolurus Wagler, 1830, Lophognathus Park. Austral Ecology 26:275-292. Gray, 1842, Rankinia Wells and Wellington, 1984, Diporiphora Broome, L. S. and Geiser, F. 1995. Hibernation in free-living Gray, 1842, Tympanocryptis Peters, 1863, as well as three new mountain pygmy-possums, Burramys parvus (Marsupialia: genera and six new subgenera. Australasian Journal of Burramyidae). Australian Journal of Zoology 43:373-379. Herpetology 30:37-64. Broome, L., Archer, M., Bates, H., Shi, H., Geiser, F., McAllan, Hoser, R. T. 2019a. 11 new species, 4 new subspecies and a B., Heinze, D., Hand, S., Evans, T. and Jackson, S. 2012. A subgenus of Australian Dragon Lizard in the genus brief review of the life history of, and threats to, Burramys parvus Tympanocryptis Peters, 1863, with a warning on the with a pre-history based proposal for ensuring that it has a conservation status and long-term survival prospects of some future. pp 114-126 in Wildlife and Climate Change: towards newly named taxa. Australasian Journal of Herpetology 39:23- robust conservation strategies for Australian fauna, Lunney, D. 52. and Hutchings, P. (eds.), Royal Zoological Society of NSW, Hoser, R. T. 2019b. Richard Shine et al. (1987), Hinrich Kaiser et Mosman, NSW, Australia. al. (2013), Jane Melville et al. (2018 and 2019): Australian Calaby, J. H., Dimpel, M. and McTaggart-Cowan, I. 1971. 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(Marsupialia: Burramyidae) and other small mammals in the Schulz, M., Wilks, G. and Broome, L. 2012b. Occupancy of spoil alpine environment at Mt Higginbotham, Victoria. Australian dumps by the Mountain Pygmy-possum Burramys parvus in Mammalogy 13:167-177. Kosciuszko National Park. Ecological Management and Mansergh, I., Heinze, D. and Weeks, A. 2010. A project to assist Restoration 13:290-296. with the conservation of the Mountain Pygmy-possum Burramys Shi, H. 2012. Thermal properties of alpine boulder fields: The parvus population on Mt Buller: facilitating and strengthening potential to provide refuge for the endangered Mountain Pygmy- genetic diversity through cross-breeding in the wild. Report to possum (Burramys parvus). PhD Thesis, University of New Department of Sustainability and Environment (Victoria). South Wales. ADFA campus ACT. Menkhorst, P. and Knight, F. 2004. A Field Guide to the Strahan, R. (ed.) 1988. 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Amphibia and Reptilia of Australia. Australian Journal of Journal of Australian Zoology 50(1):25-37. Herpetology Supplementary Series 1:1-61. Osborne, M. J., Norman, J. A., Christidis, L. and Murray, N. D. CONFLICTS OF INTEREST 2000. Genetic distinctiveness of isolated populations of an None. endangered marsupial, the mountain pygmy-possum, Burramys parvus. Molecular Ecology 9:609-613. Pepper, M., Sumner, J., Brennan, I. G., Hodges, K., Lemmon, A. R., Lemmon, E. M., Peterson, G., Rabosky, D. L., Schwarzkopf, L., Scott, I. A. W., Shea, G. and Scott keogh, J. 2018. Australasian Journal of Speciation in the mountains and dispersal by rivers: Molecularphylogeny of Eulamprus water skinks and the Herpetology ® biogeography of Eastern Australia. Journal of Biogeography. 2018;1-13. Publishes original research in printed form in relation to reptiles, other fauna and related matters, Ride, W. D. L. 1956. 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All rights reserved. scientific significance of the Australian alps. Australian Academy Australasian Journal of Herpetology is also a of Science, Canberra, ACT, Australia. registered trademark ® in all relevant areas and jurisdictions (Australian trademark number: Sanecki, G. M., Green, K., Wood, H. and Lindenmayer, D. 2006. 1686575). The implications of snow-based recreation for small mammals in the subnivean space in south-east Australia. Biological All Intellectual Property (IP) rights are reserved, Conservation 129:511-518. including in relation to all IP generated by the journal in terms of both authors, publisher and the Schulz, M., Wilks, G. and Broome, L. 2012a. An uncharacteristic like. new population of the Mountain Pygmy-possum Burramys parvus in New South Wales. Australian Zoologist 36:22-28.

Hoser 2020 - Australasian Journal of Herpetology 42:3-10. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 11 Australasian Journal of Herpetology 42:11-22. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

Small and overlooked … six new species of Pygmy Possum, Genus Cercartetus Gloger, 1841 sensu lato from the Australasian bioregion.

LSID URN:LSID:ZOOBANK.ORG:PUB:AEC58004-7BD2-4BC1-A6D4-63A29BB62FD3

RAYMOND T. HOSER LSID urn:lsid:zoobank.org:author:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 30 March 2020, Accepted 5 April 2020, Published 25 April 2020.

ABSTRACT Cercartetus Gloger, 1841, better known as the iconic Pygmy Possums are a commonly seen and familiar element of the native Australasian mammal fauna to biologists and lay people alike. While numerous forms have been described as species, as listed by Bannister et al. (1988), the taxonomy in recent years has been stable, with just four species being widely recognized. A molecular phylogeny published by Osborne and Christidis (2002) indicated that based on molecular divergences there were at least six species in the genus, five of which had available names. However as of 2019 no change to the current taxonomy or nomenclature had been formally proposed or widely adopted since at least 1934. Before formally naming the newly identified species of Osborne and Christidis (2002), all specimens of all putative species within the genus from across the known ranges of each, were examined in terms of their identification, taxonomy and nomenclature. The result was five species for which names were available and already being used (one being treated as a subspecies) and six others for which no names were available. Therefore these are formally named herein in accordance with the rules of the International Code of Zoological Nomenclature (Ride et al. 1999). Each of the six identified species (five for the first time) are morphologically divergent from their nearest congeners, reproductively isolated and geographically allopatric and separated by biogeographical barriers of known antiquity, forming a body of evidence giving a sound scientific basis for the decision to name each. With the so-called C. caudatus linage (Long-tailed Pygmy Possum) having a16-27 MYA divergence from other members of the genus (Osborne and Christidis, 2002), and morphological divergence, a strong case is made for the creation of a separate genus or subgenus for this lineage. The name Eudromicia Mjöberg, 1916 is available and therefore used. Eudromicia has been recognized in the past as a genus by Iredale and Troughton (1934). The newly named species are Cercartetus hoserae sp. nov., from north-west Victoria and nearby parts of South Australia, previously identified as a population of C. lepidus Thomas, 1888; and four others, being Eudromicia adelynhoserae sp. nov. from North Queensland, E. jackyhoserae sp. nov. from south-east Papua New Guinea (PNG), E. richardwellsi sp. nov. from the central Highlands of PNG, E. rosswellingtoni sp. nov. from the western highlands of West Papua (Irian Jaya) Indonesia and E. doriskuenae sp. nov. from high altitude areas of the Finisterre Range in Madang and Morobe Provinces, PNG, all formerly identified as populations of C. caudatus (Milne-Edwards, 1877). This paper effectively doubles the number of known extant species within the genus Cercartetus. Keywords: Mammals; Marsupial; taxonomy; nomenclature; Pygmy Possum; Cercartetus; Eudromicia; Victoria; Australia; Queensland; Irian Jaya; New Guinea; Papua; lepidus; caudatus; nanus; concinnus; macrurus; new species; hoserae; richardwellsi; adelynhoserae; rosswellingtoni; jackyhoserae; doriskuenae.

INTRODUCTION MATERIALS, METHODS AND RESULTS The iconic Pygmy Possums of the family Burramyidae Broom, Before a decision is made to name any new taxon, reasonable 1896 are a familiar element of the native Australasian mammal steps must be taken to ensure that it is justified on all relevant fauna to biologists and many lay people as well. grounds, including that it is morphologically, genetically and Cercartetus Gloger, 1841, as is widely recognized in 2019, is the reproductively isolated from their nearest relative and to a most speciose, widely distributed, abundant and best known sufficient degree to be of taxonomic significance. genus within the family. A further relevant question to ask is should the reproductively It occurs throughout the inhabited regions of southern and isolated and morphologically divergent entities be labelled as eastern Australia, including Tasmania and into New Guinea. subspecies, full species, or potentially higher level again. Where they occur, they are often abundant and are a regular Key literature relevant to the taxonomic and nomenclatural encounter in faunal field surveys and other wildlife encounters. conclusions within this paper include Anonymous (2007), Aplin While numerous forms have been described as listed by et al. (2016), Bannister et al. (1988), Bell (1828), Brammell and Bannister et al. (1988), the taxonomy in recent years has been Archer (1997), Brongersma (1953), Broom (1895, 1896a, stable, with just four species being widely recognized. 1896b), Bryant and Krosch (2016), Collins (1973), Desmarest (1818, 1820), Dwyer (1977), Eldridge et al. (2018), Flannery A molecular phylogeny published by Osborne and Christidis (1995), Gloger (1841), Gould (1845), Groves et al. (2005), Harris (2002) indicated that based on molecular divergences there (2006, 2010), Hoser (1989), Hoser (2019c), Iredale and were at least six species in the genus. Troughton (1934), Jones (1925), Krefft (1863), Lawlor (1979), Five of of these had available names. Maxwell et al. (1996), Menkhorst (2016), Menzies (1991), Milne- Their phylogeny also provided support for the contention that the Edwards (1877), Mjöberg (1916), Osborne and Christidis (2002), divergent taxon, currently known as C. caudatus (Milne- Pestell et al. (2008), Rawlings and Donnellan (2003), Ride Edwards, 1877) with a divergence estimated at a16-27 MYA (1956, 1970), Ride et al. (1999), Schäuble et al. (2000), Smith from other members of the genus (Osborne and Christidis, (1986), Strahan (1988), Thomas (1888, 1922), Wakefield (1963), 2002) could or should be placed in a separate genus. van Ufford and Cloos (2005), Vaughan (1986) and sources cited On the basis of this evidence and morphological divergence of therein (duplicitous references not necessarily included). this form and noting that it has a geographically disjunct centre Live and dead specimens as well as available bone specimens, of distribution (New Guinea, versus southern Australia), the were examined as was other relevant material, including past available name Eudromicia Mjöberg, 1916 is applied by myself climate data for the relevant regions, sea level depths, and other to the relevant species within this paper for the first time since relevant information. done so by Iredale and Troughton (1934). In summary, as inferred already, the genetic, geological, In spite of the preceding, I note that as recently as of 2019, the historical and morphological evidence clearly showed that there date this paper was prepared, that no change to the current were no less than six unnamed species in the genus. taxonomy or nomenclature of the putative genus Cercartetus SUMMARY OF THE RESULTS had been formally proposed or was being mooted by anyone. All putative recognized species of Cercartetus Gloger, 1841 Before formally naming the newly identified species of Osborne were inspected and matched with original descriptions or and Christidis (2002), all specimens of all putative species within synonymised specimens as listed by Bannister et al. (1988). the genus from across the known ranges of each, were For the relevant species the following findings are noted, along examined in terms of their identification, taxonomy and with the taxonomic implications. nomenclature including the putative new species identified by Cercartetus nanus (Desmarest, 1818), with a type locality of Osborne and Christidis (2002). Tasmania, and occurring throughout south-east Australia, As mentioned in the abstract, the final result was six species for including Tasmania has 3 available synonyms. which names were available and already being used (one Phalangista gliriformis Bell, 1828 with a type locality of Tasmania presently being treated as a subspecies) and five others for has been treated as synonymous with C. nanus and no change which no names were available. is required. Therefore these are formally named herein in accordance with Dromicia unicolour Krefft, 1863 from St. Leonards (Sydney), the rules of the International Code of Zoological Nomenclature New South Wales, has been regarded as a subspecies of C. (Ride et al. 1999). nanus. Each of the six identified species (five for the first time) are Since at least 1988, all specimens from mainland Australia have morphologically divergent from their nearest congeners, been referred to this subspecies Bannister et al. (1988). reproductively isolated and geographically allopatric and Osborne and Christidis (2002) only did a molecular analysis of separated by biogeographical barriers of known antiquity, Victorian specimens, without comparison with specimens from forming a body of evidence giving a sound scientific basis for the any parts of New South Wales, southern Queensland or decision to name each. Tasmania and so whether or not there are taxonomically The newly named species are Cercartetus hoserae sp. nov., significant divergences is not fully established. from north-west Victoria and nearby parts of South Australia, However there is no evidence of obvious morphological previously identified as a population of C. lepidus Thomas, 1888; divergence between mainland and Tasmanian specimens, or for and five others, being Eudromicia adelynhoserae sp. nov. from that matter specimens across the range on mainland Australia North Queensland, E. jackyhoserae sp. nov. from south-east and so I hereby treat Dromicia unicolour Krefft, 1863 as wholly Papua New Guinea (PNG), E. richardwellsi sp. nov. from the synonymous and not a subspecies and obviously suggest central Highlands of PNG, E. rosswellingtoni sp. nov. from the everyone else does so, unless and until compelling evidence to western highlands of West Papua (Irian Jaya) Indonesia and E. the contrary emerges. doriskuenae sp. nov. from high altitude areas of the Finisterre I note also that C. nanus remains abundant on even smaller Range in Madang and Morobe Provinces, PNG, all formerly Bass Strait Islands and these have not been formally assigned identified as populations of E. caudatus (Milne-Edwards, 1877). to either subspecies, nor do they appear to differ This paper effectively doubles the number of known extant morphologically from those on either side of the water body. species within the genus Cercartetus sensu lato. Noting that in the recent geological past (within the last 20K Within the C. caudatus species group, there may be further years), there was a wide land bridge between Tasmania and undescribed forms. Victoria, it is reasonable to expect C. nanus from either side to have mixed and without impediment, thereby maintaining

species unity. lepidus have had a long term isolation. The same biogeograpical barrier of Bass Strait has not had any Confirming this is the molecular results of Osborne and impact on most other trans-Victorian-Tasmanian species with Christidis (2002) who found species-level divergence between low habitat specificity (as is the case for C. nanus) and the two populations based on samples from each. divergence between them on either side of the now extant Bass They wrote: Strait, which is well known to have been a land bridge for most of the recent geological past. “A similar ly high le v el of DN A sequence di ver gence (c . 7%) was r ecor ded betw een the mainland and Tasmanian This includes for example Lowland Copperheads Austrelaps popula tions of C. le pidus . Although the tw o disjunct superbus (Günther, 1858) as detailed in Hoser (1989), or the populations are not currently recognised as separate frog Limnodynastes dumerilii Peters, 1863, including the subspecies, they clearly could be on the basis of DN A subspecies Limnodynastes dumerilii insularis Parker, 1940 as diver gence .” detailed by Schäuble et al. (2000) who found no significant divergence between specimens from Tasmania or nearby A sequence divergence of 7% actually implies a 3 MYA southern Victoria based on their molecular phylogeny published divergence, as in well prior to the removal of the trans-Victorian- at page 386. Tasmanian land bridge within the last 25K years and is sufficient, not just for subspecies-level recognition, but also is I note that quite significantly, the Bass Strait barrier has been species level divergence. conquered by C. nanus on both east and west sides, being found (as of now) on both King Island (west side) and Flinders A review of the extant distributions of both putative C. lepidus island (east side) and is common in all habitats on the proximal and C. nanus also yields further corroborative results beyond Victorian and Tasmanian sides of the water barrier. what I have already mentioned. On the basis of the preceding, the taxon Dromicia britta Jones, Based on Museum specimen records of all relevant Australian 1925 with a type locality of Millicent, South Australia is also State Museums, downloaded via the “Atlas of Living Australia” herein treated as a synonym of C. nanus and not putative C. (ALA) at “https://www.ala.org.au/” (last downloaded on 10 March nanus unicolour. 2020), one finds that C. nanus remains common on both Flinders and King Islands in Bass Strait, implying that at the time The species Cercartetus concinnus (Gould, 1845), type locality of glacial maxima, this was the taxon that inhabited this region Swan River (Perth) Western Australia, was redescribed by (Bass Strait) and noting that it tends to live exclusive of putative Waterhouse in 1846 as Phalangista (Dromica) neillii based on C. lepidus, this in effect confirms the long historical gap between another specimen from nearby King George Sound in Western the two populations of this putative species. Australia. Wakefield (1963) named a specimen from Natimuk in Victoria as By contrast (and based on ALA records), putative C. lepidus is a subspecies Cercartetus concinnus minor. However Osborne not found on King Island, being the western and logical land and Christidis (2002) Wrote: bridge to its nearest point on the mainland side of the Bass Strait. Putative C. lepidus is found on Flinders Island on the east “Flannery (1994) recognised two subspecies of C. concinnus, C. side of Bass Strait, but the nearest mainland point to here c. concinnus in southwest Western Australia, and C. c. minor in Wilsons Promontory, is a stronghold for C. nanus, as are all South Australia, western Victoria and western New South Wales. areas anywhere within 1,000 km of here (as measured along the Conversely, Strahan (1995) did not recognise any subspecies. extant Victorian coastline), in effect preventing the Tasmanian The low level of divergence that was recorded between putative C. lepidus from having any contact with the mainland individuals of C. concinnus from western Australia and south- population. western New South Wales supports the latter view. There was less than 1% divergence between the two disjunct populations, Coupled with the fact that the mainland specimens of C. lepidus which was only slightly higher than that which was identified have significantly divergent morphology and biology, the case for between two individuals from the same locality (Toodyay, formally recognizing them as a new species is compelling. Western Australia). They are herein identified as C. hoserae sp. nov.. On the basis of the preceding I agree that there is no point in It is also worth noting that there is a similar divergence to the recognising the eastern population of C. concinnus as a above Cercartetus species seen between the agamid species subspecies. Rankinia diemensis (Gray, 1841) from Tasmania and R. neildaviei Hoser, 2015 also restricted to a region in western The taxonomic situation with respect to the other two putative Victoria (Hoser, 2015) and that both diverged well prior to the species of Cercartetus is however very different and needs to be presence of a land bridge across the Bass Strait, indicating a elaborated upon to explain the scientific basis for the taxonomic non-marine barrier between the two evolving species. judgements in this paper. The putative species C. caudatus (Milne-Edwards, 1877) was The putative species Cercertatus lepidus (Thomas, 1888) with a described from a specimen in the Arfak Mountains in Irian Jaya, type locality of Tasmania is known from most parts of Tasmania which is particularly significant in light of what follows. as well as region in Western Victoria, bounded by the Little Desert in the South, the Murray River in the north and nearby Another putative taxon, “Eudromicia macrura Mjöberg, 1916”, drier parts of south-east South Australia although there is recent based on a specimen from Cedar Creek, Atherton, Queensland, (aged) fossil material from nearby on the north (NSW) side of Australia (see comments on Eudromicia below) has since been Murray River (Strachan 1988). synonymised with C. caudatus as seen for example in Osborne and Christidis (2002) at fig. 5. In the wild, this species appears not to be found in areas where Other authors, including Strahan (1988) at page 166 refer to the C. nanus occurs, this especially being the case on the mainland more recently described form as a subspecies, namely C. of Australia, including far southern Victoria, where C. nanus is caudatus macrurus, but fail to give any morphological basis of strong and there are no putative C. lepidus at all. separation. On the mainland of south-east Australia where both C. nanus and putative C. lepidus occur, C. nanus is found in wetter Significantly Osborne and Christidis (2002) tested samples of regions (such as southern Victoria) and C. lepidus in drier areas the putative species from both New Guinea and Australia and further north. This in effect separates the populations of C. found significant divergence between the two. lepidus in Tasmania and putative C. lepidus from mainland They wrote: Australia, even without consideration of the trans-Victorian- “signif icant genetic differentia tion w as obser ved within Tasmanian barrier. some species. Most di ver ged (8%) w ere C. cauda tus macrurus and C. cauda tus cauda tus fr om A ustr alia and On its own this implies that the two populations of putative C.

Hoser 2020 - Australasian Journal of Herpetology 42:11-22. N e w Guinea r especti vely. ... T he mainland and Tasmanian Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 14 Australasian Journal of Herpetology

popula tions of C. le pidus and the A ustr alian and Ne w altitude barriers in times of glacial maxima has been due to a Guinean popula tions of C. cauda tus ar e estima ted to have parallel drying of climates thereby in effect shrinking rainforest diverged c . 3-4 million y ears ago.” areas and in fact potentially further confining the relevant On face value these results indicated that elevation of the species. putative subspecies C. caudatus macrurus to full species was Also not considered by many zoologists is the underlying fact an obvious decision to make. that in times of glacial maxima, the cooling is most extreme in However inspection of specimens used for analysis and the the polar and near polar regions and reduces towards the paper of Osborne and Christidis (2002) raised further important equatorial regions, lessening the impact of global cooling. issues. This means temperature drops near the equator are not The putative C. caudatus material from New Guinea that was sufficient to enable a significant downward elevation migration of tested by Osborne and Christidis (2002) was not taken from the species confined to so called sky islands. type locality on the far west of the island, but instead came from Noting that molecular studies on similarly constrained species of a geographically distant region in the south-east of New Guinea reptiles (as cited by Hoser 2016), Moritz et al. (1993) and at Milne Bay Province from an Australian Museum specimen mammals, such as Tree Kangaroos (Dendrolagus Müller, 1840 number 24767. species) as detailed by Eldridge et al. (2018) and Hoser (2019) This fact necessitated an inspection of specimens from this came to the same results. That is a well-established trend of locality to compare them with others from the type locality of the cold-climate, rainforest obligate species in north-east Arfak Mountains in Irian Jaya, which is far west of the main Queensland and New Guinea being unable to cross even central cordillera of the island of Papua and therefore in a region relatively short distances across unsuitable lowland dry zones. well and truly separated by unsuitable habitat, as well as It is therefore self evident that even before a morphological specimens from intermediate locations elsewhere on the island study is completed, that the Milne Bay putative C. caudatus of New Guinea to ascertain whether they fitted within one or both must be a separate taxon to those from the Arfak Mountains putative species, or alternatively were different taxa yet again. specimens. Putative C. caudatus in Australia, where they have been well Collection of putative C. caudatus in New Guinea has shown the studied are very habitat restricted and range restricted, which putative species to be abundant where it occurs, but this also raised issues in terms of Australian specimens and whether abundance is heavily restricted to well-known upland areas that or not more than one species is involved. are in turn separated by lower altitude regions where the In terms of all known populations and summarizing current putative species appears absent. knowledge, Aplin et al. (2008) wrote of putative C. caudatus: The clustering of records is so significant as compared to other “This species is present in the highlands of the island of New New Guinea vertebrates in Museum collections that one can Guinea (Indonesia and Papua New Guinea), and in the lowland only conclude that the absence in intervening regions is due to and upland rainforests between Paluma Range and Cooktown, absence of specimens and not absence of collecting. Queensland, Australia (Flannery 1995, Maxwell et al. 1996). It Hence, with at least five separate populations of putative C. ranges from medium elevations to 3,450 m asl (New Guinea) caudatus on the island of New Guinea, clearly separated by and from sea level to 1,600 m in Australia. ... currently unpassable altitudinal and habitat barriers (and other In New Guinea, the species is found in montane and mid- subpopulations that may also be taxonomically distinct), it montane primary and secondary tropical moist forests, seems self evident that each would require taxonomic especially where there are tree ferns. It also occurs in areas of recognition if each were clearly morphological divergent. subalpine shrubland. In Australia, it is known from rainforest This in fact is the case and exactly why I have no hesitation in (Maxwell et al. 1996).” describing four obviously divergent high-altitude forms from New Significantly and in error, Aplin et al. (2008) published a Guinea, separated by well known biogeographical barriers as distribution map for putative C. caudatus that showed a new species. continuous line of distribution along the centre of the island from Plotting the New Guinea distribution of putative C. caudatus the birds head in the far north-west, across the low-lying area based on ALA records, one finds a near identical distribution joining this part of the island to the rest and then continuously pattern to that of New Guinea Tree Kangaroos (Dendrolagus) in along the central cordillera to the Milne Bay area in the far the so-called D. dorianus Ramsay, 1883 species-group as south-east. shown in Eldridge et al. (2018) on page 590. The ALA records (last downloaded on 10 March 2020), and Pages 593 and 594 of Eldridge et al. (2018) show clear species records on Vert Net (VN) at http://vertnet.org/ (last downloaded level divergences of the Dendrolagus species constrained by the on 10 March 2020), this being an international database of same biogeographical factors as the putative C. caudatus, museum collections, shows the distribution of putative C. making species level recognition the logical outcome of this caudatus to in fact be restricted to a small number of isolated analysis. high altitude pockets and a complete absence in intervening In the Australian context, the results of Eldridge et al. (2018) can areas. literally be overlain to putative C. caudatus as a direct pointer to Between the Arfak Mountains in Irian Jaya on the so-called the likelihood of there being two species in the Australian wet “Bird’s head” of the island, and the main central cordillera is a tropics as opposed to just one. Again refer to several other wide region of flat land or of generally low hills separating these examples in Hoser (2016) and Moritz et al. (1993). high elevation biogeographical features effectively separating A check of the ALA shows the putative species is constrained in obligate highland rainforest-type species. a similar way to the two recognized species of Dendrolagus While periods of glacial maxima and significantly cooler global across the same barrier just north of Cairns, Australia, in this climates could be interpreted as lowering altitudinal limits and case being estimated at about 2 MYA. Hence a similar allowing species to cross these barriers and this is what many divergence between the two disjunct Australian populations of zoologists have assumed, evidence on all similarly constrained putative C. caudatus can be readily accepted as fact and in the species groups has shown that in recent glacial minima this absence of specific DNA testing of specimens from both sides effect has not happened! of the barrier. See for example the cases documented by Hoser (2016). The geoclimatic evidence is irrefutable. The overlooked reason for the apparent inability of these cool Significantly, specimens from the north of the biogeograhical climate rainforest obligate species to be able to cross the low gap north of Cairns, sometimes called the “Black Mountain Gap”

(Bryant and Krosch 2016) are morphologically divergent from zoologists (three other long recognized species), is effectively those to the south and hence I also have no hesitation in left untouched, save for the formal naming of a well-known and formally naming this taxon as a new species. divergent form, giving an effective 7/4 ratio for the number of An outlier population of putative C. caudatus from high altitude species in the putative genus. areas of the Finisterre Range in Madang and Morobe Provinces, However the morphological and genetic evidence supports this PNG, separated from the nearby Central Highlands of PNG by position. the Ramu and Markham River basins is also sufficiently I also note that the phylogeny published by Osborne and divergent as to warrant full species recognition. Christidis (2002) on page 32 shows the putative taxon C. In summary, C. caudatus as currently recognized is split into a caudatus as basal to the rest, thereby enabling a greater total of seven separate species. This is two from Australia and potential time frame to speciate (all other factors being equal) five from New Guinea. and note that two geographically proximal samples from across With a divergence time estimated at 16-27 MYA for the C. the range of the putative taxon did in fact show species-level caudatus linage versus the rest of the genus (Osborne and divergence for these populations in their results. This also Christidis, 2002), a strong case is made for the creation of a implies a potentially even greater divergence for geographically separate genus or subgenus for this lineage. distant samples. It is also significant that the habitat fidelity and dispersal abilities Adding morphological divergence, the case for genus level of the species in the genus Eudromicia are significantly and recognition of the species group becomes compelling. demonstrably lower than for Cercartetus as demonstrated by the The name Eudromicia Mjöberg, 1916 is available and has been date of divergence of all species in the former genus across recognized in the past as a genus by Iredale and Troughton known biogeographical barriers, versus those for Cercartetus. (1934) and is a contention I agree with. Again this aids the contention that there has in fact been more Hence the relevant new species are herein placed within this speciation and over a longer time frame for the genus genus and not Cercartetus sensu stricto. Eudromicia as opposed to Cercartetus. The newly named Australian form is that from the highlands of INFORMATION RELEVANT TO THE FORMAL DESCRIPTIONS the northern wet tropics, Queensland, centred on Mount Lewis THAT FOLLOW and it is formally named Eudromicia richardwellsi sp. nov. There is no conflict of interest in terms of this paper or the Within the island of New Guinea, the species E. caudatus is conclusions arrived at herein. restricted to the Arfak Mountains in Irian Jaya on the so-called Several people including anonymous peer reviewers who “Bird’s Head” of the island. revised the manuscript prior to publication are also thanked as The species E. adelynhoserae sp. nov. is the taxon from the are relevant staff at museums who made specimens and Milne Bay region of Papua New Guinea, tentatively including the records available in line with international obligations. population from offshore Fergusson Island (max. elevation is In terms of the following formal descriptions, spellings should 1947 m above sea level), based on proximity to adjacent not be altered in any way for any purpose unless expressly and mainland populations and similarity of reptile taxa between the exclusively called for by the rules governing Zoological two places (e.g. the python Lenhoserus boeleni (Brongersma, Nomenclature as administered by the International Commission 1953)). of Zoological Nomenclature. The species E. rosswellingtoni sp. nov. is the taxon from the In the unlikely event two or more newly named taxa are deemed central Highlands of PNG, including the nearby high altitude conspecific by a first reviser, then the name to be used and points in the Eastern Highlands province and Southern retained is that which first appears in this paper by way of page Highlands Province. priority and as listed in the abstract keywords. The species E. jackyhoserae sp. nov. is the taxon from the Some material in descriptions for taxa may be repeated for other Nassau Range in Irian Jaya (West Papua). taxa in this paper and this is necessary to ensure each fully The species E. doriskuenae sp. nov. is the taxon from the high complies with the provisions of the International Code of altitude areas of the Finisterre Range in Madang and Morobe Zoological Nomenclature (Fourth edition) (Ride et al. 1999) as Provinces, PNG. amended online since. One or more further potentially unnamed species may be Material downloaded from the internet and cited anywhere in this present from Foja Mountains north of the main cordillera in Irian paper was downloaded and checked most recently as of 10 Jaya (near the coast) as detailed by Anonymous (2007) (and March 2020 (including if also viewed prior), unless otherwise see image at: https://www.naturepl.com/stock-photo-long-tailed- stated and was accurate in terms of the content cited herein as pygmy-possum-nature-image01361572.html) (Last downloaded of that date. on 10 March 2020), but I have deferred formally naming the Unless otherwise stated explicitly, colour and other descriptions taxon on the basis that it is morphologically similar to the apply to living adult specimens of generally good health and not specimens from the mid-central highlands and because of the under any form of stress by means such as excessive cool, recent geological age of the highlands where it occurs. heat, dehydration or abnormal skin reaction to chemical or other The preceding would mean an estimated divergence from the input. central highlands stock of about 1.25 MYA, which at best would Colour descriptions of species refer to fur colour and not skin. usually be defined as a subspecies, rather than a full species. While numerous texts and references were consulted prior to Also refer to the Dendrolagus phylogeny published by Eldridge publication of this paper, the criteria used to separate the et al. (2018). relevant species or subspecies has already been spelt out and/ Of course there is also potential for one or more other unnamed or is done so within each formal description and does not rely on forms on the island of New Guinea, particularly in isolated high material within publications not explicitly cited herein. regions of significant antiquity, but so far there is no indication as In terms of conservation of each population of each species as to exactly where these may occur. described below, the comments in Hoser (1991 and 2019a, It is noted that a person unfamiliar with the relevant taxa and not 2019b) apply. reading the detail of this paper and the sources cited herein, Wildlife laws as currently enforced in Australia, Indonesia and may form an initial view that the “splitting” of one putative Papua New Guinea or not in a materially significant way species to seven is unjustified, while the remainder of the genus enhancing the long-term survival prospects of any of the Cercartetus sensu lato as currently understood by most relevant species.

Over breeding of humans and the environmental problems adelynhoserae sp. nov. (from Milne Bay, PNG), E. associated with this overpopulation are by far the greatest long rosswellingtoni sp. nov. (from the central ranges of New Guinea, term threat to each and every relevant species. generally near Mount Wilhelm to the Irian Jaya border) and E. In line with the Australian Federal Government’s “Big Australia” jackyhoserae sp. nov. (from Nassau Range, Irian Jaya) are policy, that being to increase the human population of 25 million readily separated from all other members of the genus by having (2020), from 13 million in around 1970, to over 100 million within a large size, as in being more than 85 mm head and body 100 years “so that we can tell China what to do”, as stated by length; lower leg length more than 20 mm long and hind leg the former Prime Minister, Kevin Rudd in 2019 (Zaczek 2019), more than 13 mm long. The tail is also long and more than 120 the human pressure on the relevant ecosystems has increased mm in length, slightly thickened and furred at the base in line with the human populations nearby and will clearly (separating it from the superficially similar Pogonomys Milne- continue to do so. Edwards, 1877). The dark mark through the eye is obvious and CERCARTETUS HOSERAE SP. NOV. there are large crinkly ears. This is also a genus-level diagnosis for all species within Eudromicia. LSID urn:lsid:zoobank.org:act:3AFEEC0A-8AFB-4E49-B254- 9298BC427BA7 Other characteristics diagnostic of Eudromicia include large eyes, mouse-like ears, a pouch that opens anteriorly (as in Holotype: A preserved male specimen at the National Museum didelphids), a well-developed and opposable hallux that does not of Victoria, Melbourne, Victoria, Australia, specimen number have a claw, expanded pads at the ends of the digits, C27321 collected from 6.6 km west of Sunset Tank, Victoria, quadritubercular molars, a broad, flattened skull, and three Australia, Latitude -34.95 S., Longitude 141.45 E. This upper incisors and one lower incisor that are enlarged and government-owned facility allows access to its holdings. procumbent. (Lawlor 1979, Ride 1970, Strahan 1988, Vaughan Paratype: A preserved male specimen at the National Museum 1986). of Victoria, Melbourne, Victoria, Australia, specimen number Distribution: Cercartetus hoserae sp. nov. is restricted to a dry C24355 collected at 23 km south of Tutye, Victoria, Australia, zone of habitats in Western Victoria, from Little Desert in the Latitude -35.42 S., Longitude 141.48 E. south to the Murray River in the north and west into nearby parts Diagnosis: The genus Cercartetus Gloger, 1841 is effectively of South Australia. The range is bounded by the Murray River diagnosed by Thomas (1888) at pages 140-141 under the name drainage to the north and west, wetter habitats to the south, Dromicia Gray, 1841. where C. nanus occurs and agricultural regions further east in The putative species Cercartetus lepidus (Thomas, 1888) Victoria. including the species C. hoserae sp. nov. (from Victoria and C. lepidus is confined to Tasmania and is therefore an endemic South Australia) and C. lepidus (from Tasmania) are both to that state. separated from all other species in the genus (sensu lato and Etymology: Named in honour of my wife, Shireen Vanessa including the genus Eudromicia Mjöberg, 1916 as defined below) Hoser, of Park Orchards, Victoria, Australia, but born in a remote by the following unique suite of characters: Having slate part of southern Africa called “Athlone”, not far from central coloured belly hairs basally and of a small size, being less than Cape Town in recognition of her services to herpetology and 85 mm head and body length; lower leg less than 20 mm long wildlife conservation over more than 20 years. and hind leg less than 13 mm long as well as a fourth small molar (the final character which in particular separates this EUDROMICIA RICHARDWELLSI SP. NOV. species from C. nanus (Desmarest, 1818) and C. concinnus LSID urn:lsid:zoobank.org:act:92B808AB-709C-4FDC-89CA- (Gould, 1865)). 4728900E986E C. hoserae sp. nov. is readily separated from C. lepidus by Holotype: A preserved female specimen in the Queensland having brownish yellow-fur on the upper body, versus grey with a Museum, Brisbane, Queensland, Australia, specimen number slight brown tinge or if yellowish, with a strong grey infusion of JM7011, collected from Mount Spurgeon, Queensland, Australia, colour in C. lepidus. C. hoserae sp. nov. is further separated Latitude -16.6 S., Longitude 145.2 E. This government-owned from C. lepidus by having an obvious demarcation between the facility allows access to its holdings. blackish fur around the eye and the pink of the snout, this being Paratypes: Three preserved specimens in the Queensland a line of brownish yellow-fur, whereas in C. lepidus blackish fur Museum, Brisbane, Queensland, Australia, specimen numbers runs in some form to the snout. JM10205, JM10206 and JM10228 (skulls), collected from Mount A detailed morphological appraisal of the holotype of C. lepidus, Lewis, Queensland, Australia, Latitude -16.58 S., Longitude also applicable to C. hoserae sp. nov. can be found on pages 145.28 E. 142 and 143 of Thomas (1888). Diagnosis: The species complex, until now regarded as The species Cercartetus nanus (Desmarest, 1818) is separated putative Eudromicia (formerly Cercartetus) caudatus (Milne- from all other species in the genus by having a medium tail Edwards, 1877) (herein restricted to the type region of the type being less than 120 mm in length; an indistinct darker eye-mark; locality, being the Arfak Mountains in western Papua, ventral hairs tipped with white (versus completely white in C. Indonesia), including E. macrurus (Mjöberg, 1816) (from the concinnus (Gould, 1865)). southern wet tropics in Queensland, Australia) and the four The species C. concinnus (Gould, 1865) is separated from all newly described species, all formerly treated as populations of other species in the genus by having a small size as for C. E. caudatus, namely E. richardwellsi sp. nov. (from the northern lepidus and C. hoserae sp. nov. but is easily separated from wet tropics, Queensland Australia), E. adelynhoserae sp. nov. those species by having pure white hairs on the belly and an (from Milne Bay, PNG), E. rosswellingtoni sp. nov. (from the almost obsolete (indistinct) eye mark; three molar teeth on each central ranges of New Guinea, generally near Mount Wilhelm to side, top and bottom, with the upper third premolar with a single the Irian Jaya border), E. jackyhoserae sp. nov. (from Nassau point and the lower third premolar being tiny. Range, Irian Jaya) and E. doriskuenae sp. nov. from the high altitude areas of the Finisterre Range in Madang and Morobe The species complex, until now regarded as putative Provinces, PNG, are readily separated from all other members Eudromicia, formerly Cercartetus caudatus (Milne-Edwards, of the genus by having a large size as in being more than 85 1877) (from western Papua), including E. macrurus (Mjöberg, mm head and body length; lower leg length more than 20 mm 1816) (from the southern wet tropics in Queensland, Australia) long and hind leg more than 13 mm long. The tail is also long and the four newly described species, all formerly treated as and more than 120 mm in length, slightly thickened and furred at populations of E. caudatus, namely E. richardwellsi sp. nov. the base (separating it from the superficially similar Pogonomys (from the northern wet tropics, Queensland Australia), E. Milne-Edwards, 1877). The dark mark through the eye is obvious

and there are large crinkly ears. This is also a genus-level The belly is beige and the lower flanks have well defined diagnosis for all species within Eudromicia. demarcation between darker upper parts and lighter underparts, Eudromicia caudatus (Milne-Edwards, 1877) herein restricted to which do not run in an even line. The upper limbs and lower the type region of the type locality, being the Arfak Mountains in flanks are dark brown with well defined lighter edges, forming western Papua, Indonesia and all other species in the complex, large patches and quite distinct from the brownish grey dorsal are easily separated from one another on colouration. surfaces. Eudromicia caudatus (Milne-Edwards, 1877) is readily separated The species E. doriskuenae sp. nov. from the high altitude areas from the other five herein recognized species by having a dull of the Finisterre Range in Madang and Morobe Provinces, PNG rufous colour above. The face is rufous fawn, with two broad is similar in most respects to E. rosswellingtoni sp. nov. however black bands passing from the nose through the eyes, but not can be differentiated from that species and in turn all others, by quite reaching the ears. Ears are rather long and narrow, evenly the fact that demarcation between darker upper parts and lighter oval, naked. Back-hairs are slaty grey for five sixths of their underparts is not well defined; inner ears are orange in colour length, their tips dull rufous. Chest and belly pale yellowish except for the very outer tips which have grey outer edges; white, the bases of the hairs slate-coloured. around the eye, the black marking is dark brown rather than Arms and legs grey; hands and feet dull fawn. black with some infusions of brown into the dark brown region anterior to the eye. E. macrurus (Mjöberg, 1816) from the southern wet tropics in Eudromicia macrurus (Mjöberg, 1816) in life are seen in Strahan Queensland, Australia, generally south and west of Cairns, (1998) on page 167 (all photos) and also online at: North Queensland are readily separated from the other five herein recognized species by having a brownish-grey colour https://www.flickr.com/photos/euprepiosaur/8548752100/ above, with a slight rufous tinge at the anterior end only. The Eudromicia hoserae sp. nov. in life can be seen in a photo online face is rufous fawn or just yellowish brown, depending on the at: individual, with a square black patch across each eye and https://www.flickr.com/photos/mattsummerville/43996071634/ nowhere near the ears. This patch is notable for having a well Distribution: E. richardwellsi sp. nov. occurs in the northern wet defined black triangle jutting out from above the rear of the eye, tropics of Australia from north of Port Douglas and south of the apex pointing towards the crown. Cape Tribulation, with a distribution centred on the Mount Lewis Ears are rather long and narrow, evenly oval, naked and greyish area, Queensland, Australia, being restricted to pockets of on the lobes and rich pink within. Back-hairs are slaty grey for rainforest habitat. five sixths of their length, their tips a brownish-grey colour. Chest Etymology: Named in honour of Richard W. Wells, currently of and belly pale yellowish white, the bases of the hairs slate- Lismore, New South Wales, Australia in recognition of his coloured. significant taxonomic works on Australian herpetology including Arms and legs are brown on top, light (whitish) underneath; major monographs in the 1980’s (e.g. Wells and Wellington hands and feet a light pinkish colour. 1984, 1985) and the post year 2000 period. E. richardwellsi sp. nov. from the northern wet tropics generally EUDROMICIA ADELYNHOSERAE SP. NOV. from north of Port Douglas and south of Cape Tribulation, with a LSID urn:lsid:zoobank.org:act:1102DEB8-FD63-41F0-B4E1- distribution centred on the Mount Lewis area is similar in most EDF0C0C1BCAC respects to E. macrurus, which it would otherwise be identified as. E. richardwellsi sp. nov. can however be separated from E. Holotype: A preserved specimen at the Australian Museum in macrurus by having significantly less-well defined black around Sydney, New South Wales, Australia, specimen number the eye, being peppered in appearance and not boldly M.29554, collected from Param village gardens, Aguan Area, demarcated from the adjacent brownish fur as seen in E. Milne Bay Province, Papua New Guinea, Latitude -9.97 S., macrurus. The jowls region in E. richardwellsi sp. nov. is yellow Longitude 149.48 E. This government-owned facility allows rather than whitish as seen in E. macrurus. access to its holdings. The species E. adelynhoserae sp. nov. from the Milne Bay Paratypes: Two preserved specimens at the Australian Museum Province, Papua New Guinea is readily separated from the other in Sydney, New South Wales, Australia, specimen numbers species by having a distinctly steel-greyish colour, contrasting M.29555 and M.29553 collected from Param village gardens, with white undersides and jowls. The upper surface has a Aguan Area, Milne Bay Province, Papua New Guinea, Latitude - somewhat peppered appearance. The ears are blackish grey, 9.97 S., Longitude 149.48 E. the dark colour extending almost all the way into the visible ear, Diagnosis: The species complex, until now regarded as with inner regions a pale orangeish colour. The black patches putative Eudromicia (formerly Cercartetus) caudatus (Milne- over the eyes are elliptical in shape and extend from the snout. Edwards, 1877) (herein restricted to the type region of the type There is an ill defined line of blackish grey running from the mid- locality, being the Arfak Mountains in western Papua, snout between the eyes, further back to the crown. Indonesia), including E. macrurus (Mjöberg, 1816) (from the The species E. rosswellingtoni sp. nov. from the region around southern wet tropics in Queensland, Australia) and the four Mount Wilhelm and including nearby elevated parts of Eastern newly described species, all formerly treated as populations of Highlands and Southern Highlands provinces of Papua New E. caudatus, namely E. richardwellsi sp. nov. (from the northern Guinea, west to the Irian Jaya border is readily separated from wet tropics, Queensland Australia), E. adelynhoserae sp. nov. the other species by having brown hair on the upper body, ears (from Milne Bay, PNG), E. rosswellingtoni sp. nov. (from the that are brownish grey at the outer edges and becoming orange central ranges of New Guinea, generally near Mount Wilhelm, further in, feet are pink and the pink of the snout almost extends west to near the Irian Jaya border), E. jackyhoserae sp. nov. to the eyes. The belly is beige and the lower flanks have well (from Nassau Range, Irian Jaya) and E. doriskuenae sp. nov. defined demarcation between darker upper parts and lighter from the high altitude areas of the Finisterre Range in Madang underparts, which do not run in an even line. and Morobe Provinces, PNG, are readily separated from all The species E. jackyhoserae sp. nov. from the Nassau Range in other members of the genus by having a large size as in being Irian Jaya (West Papua), Indonesia is similar in most respects to more than 85 mm head and body length; lower leg length more E. rosswellingtoni sp. nov. as described above, but can be than 20 mm long and hind leg more than 13 mm long. The tail is separated from that species by having orange-pink feet and also long and more than 120 mm in length, slightly thickened mottled darker patches or blotches on the sides of the face and and furred at the base (separating it from the superficially similar forelimbs. The ears have greyish tips, but are otherwise Pogonomys Milne-Edwards, 1877). The dark mark through the yellowish in colour further in. eye is obvious and there are large crinkly ears. This is also a

Hoser 2020 - Australasian Journal of Herpetology 42:11-22. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 18 Australasian Journal of Herpetology genus-level diagnosis for all species within Eudromicia. demarcation between darker upper parts and lighter underparts, Eudromicia caudatus (Milne-Edwards, 1877) herein restricted to which do not run in an even line. The upper limbs and lower the type region of the type locality, being the Arfak Mountains in flanks are dark brown with well defined lighter edges, forming western Papua, Indonesia and all other species in the complex, large patches and quite distinct from the brownish grey dorsal are easily separated from one another on colouration. surfaces. Eudromicia caudatus (Milne-Edwards, 1877) is readily separated The species E. doriskuenae sp. nov. from the high altitude areas from the other five herein recognized species by having a dull of the Finisterre Range in Madang and Morobe Provinces, PNG rufous colour above. The face is rufous fawn, with two broad is similar in most respects to E. rosswellingtoni sp. nov. however black bands passing from the nose through the eyes, but not can be differentiated from that species and in turn all others, by quite reaching the ears. Ears are rather long and narrow, evenly the fact that demarcation between darker upper parts and lighter oval, naked. Back-hairs are slaty grey for five sixths of their underparts is not well defined; inner ears are orange in colour length, their tips dull rufous. Chest and belly pale yellowish except for the very outer tips which have grey outer edges; white, the bases of the hairs slate-coloured. around the eye, the black marking is dark brown rather than Arms and legs grey; hands and feet dull fawn. black with some infusions of brown into the dark brown region anterior to the eye. E. macrurus (Mjöberg, 1816) from the southern wet tropics in Distribution: E. adelynhoserae sp. nov. is known from elevated Queensland, Australia, generally south and west of Cairns, rainforest locations in the Milne Bay province of the island of North Queensland are readily separated from the other five New Guinea, generally near the type locality. Fergusson Island herein recognized species by having a brownish-grey colour specimens of this genus are presumably of the same species. above, with a slight rufous tinge at the anterior end only. The face is rufous fawn or just yellowish brown, depending on the Etymology: Named in honour of my eldest daughter, Adelyn individual, with a square black patch across each eye and Hoser of Park Orchards, (Melbourne), Victoria, Australia in nowhere near the ears. This patch is notable for having a well recognition of over 20 years of significant contributions to wildlife defined black triangle jutting out from above the rear of the eye, conservation in Australia, including her work with Snakebusters: the apex pointing towards the crown. Australia’s best reptiles shows, being the only hands on wildlife Ears are rather long and narrow, evenly oval, naked and greyish shows in Australia that let people hold the animals. on the lobes and rich pink within. Back-hairs are slaty grey for Adelyn Hoser also made significant contributions to several five sixths of their length, their tips a brownish-grey colour. Chest years research on the Mountain Pygmy Possums of the genus and belly pale yellowish white, the bases of the hairs slate- Burramys Broom (1896) in the snowfields of New South Wales coloured. and Victoria, Australia. Arms and legs brown on top, light (whitish) underneath; hands EUDROMICIA ROSSWELLINGTONI SP. NOV. and feet a light pinkish colour. LSID urn:lsid:zoobank.org:act:186515F3-480A-4B09-9EA8- 47C19A3152F5 E. richardwellsi sp. nov. from the northern wet tropics generally from north of Port Douglas and south of Cape Tribulation, with a Holotype: A preserved specimen at the Australian Museum in distribution centred on the Mount Lewis area is similar in most Sydney, New South Wales, Australia, specimen number M.9622 respects to E. macrurus, which it would otherwise be identified collected from Korn in the Upper Waghi Valley, Western as. E. richardwellsi sp. nov. can however be separated from E. Highlands Province, New Guinea, Latitude -5.83 S., Longitude macrurus by having significantly less-well defined black around 144.32 E. This facility allows access to its holdings. the eye, being peppered in appearance and not boldly Paratypes: Two preserved specimens at the Australian Museum demarcated from the adjacent brownish fur as seen in E. in Sydney, New South Wales, Australia, specimen numbers macrurus. The jowls region in E. richardwellsi sp. nov. is yellow M.9621 and M.9620 collected from Korn in the Upper Waghi rather than whitish in E. macrurus. Valley, Western Highlands Province, New Guinea, Latitude -5.83 The species E. adelynhoserae sp. nov. from the Milne Bay S., Longitude 144.32 E. Province, Papua New Guinea is readily separated from the other Diagnosis: The species complex, until now regarded as species by having a distinctly steel-greyish colour, contrasting putative Eudromicia (formerly Cercartetus) caudatus (Milne- with white undersides and jowls. The upper surface has a Edwards, 1877) (herein restricted to the type region of the type somewhat peppered appearance. The ears are blackish grey, locality, being the Arfak Mountains in western Papua, the dark colour extending almost all the way into the visible ear, Indonesia), including E. macrurus (Mjöberg, 1816) (from the with inner regions a pale orangeish colour. The black patches southern wet tropics in Queensland, Australia) and the four over the eyes are elliptical in shape and extend from the snout. newly described species, all formerly treated as populations of There is an ill defined line of blackish grey running from the mid- E. caudatus, namely E. richardwellsi sp. nov. (from the northern snout between the eyes, further back to the crown. wet tropics, Queensland Australia), E. adelynhoserae sp. nov. The species E. rosswellingtoni sp. nov. from the region around (from Milne Bay, PNG), E. rosswellingtoni sp. nov. (from the Mount Wilhelm and including nearby elevated parts of Eastern central ranges of New Guinea, generally near Mount Wilhelm Highlands and Southern Highlands provinces of Papua New and west to the Irian Jaya border), E. jackyhoserae sp. nov. Guinea and west to the Irian Jaya border is readily separated (from Nassau Range, Irian Jaya) and E. doriskuenae sp. nov. from the other species by having brown hair on the upper body, from the high altitude areas of the Finisterre Range in Madang ears that are brownish grey at the outer edges and becoming and Morobe Provinces, PNG, are readily separated from all orange further in, feet are pink and the pink of the snout almost other members of the genus by having a large size as in being extends to the eyes. The belly is beige and the lower flanks have more than 85 mm head and body length; lower leg length more well defined demarcation between darker upper parts and lighter than 20 mm long and hind leg more than 13 mm long. The tail is underparts, which do not run in an even line. also long and more than 120 mm in length, slightly thickened The species E. jackyhoserae sp. nov. from the Nassau Range in and furred at the base (separating it from the superficially similar Irian Jaya (West Papua), Indonesia is similar in most respects to Pogonomys Milne-Edwards, 1877). The dark mark through the E. rosswellingtoni sp. nov. as described above, but can be eye is obvious and there are large crinkly ears. This is also a separated from that species by having orange-pink feet and genus-level diagnosis for all species within Eudromicia. mottled darker patches or blotches on the sides of the face and Eudromicia caudatus (Milne-Edwards, 1877) herein restricted to forelimbs. The ears have greyish tips, but are otherwise the type region of the type locality, being the Arfak Mountains in yellowish in colour further in. western Papua, Indonesia and all other species in the complex, The belly is beige and the lower flanks have well defined are easily separated from one another on colouration.

Eudromicia caudatus (Milne-Edwards, 1877) is readily separated The species E. doriskuenae sp. nov. from the high altitude areas from the other five herein recognized species by having a dull of the Finisterre Range in Madang and Morobe Provinces, PNG rufous colour above. The face is rufous fawn, with two broad is similar in most respects to E. rosswellingtoni sp. nov. however black bands passing from the nose through the eyes, but not can be differentiated from that species and in turn all others, by quite reaching the ears. Ears are rather long and narrow, evenly the fact that demarcation between darker upper parts and lighter oval, naked. Back-hairs are slaty grey for five sixths of their underparts is not well defined; inner ears are orange in colour length, their tips dull rufous. Chest and belly pale yellowish except for the very outer tips which have grey outer edges; white, the bases of the hairs slate-coloured. around the eye, the black marking is dark brown rather than Arms and legs grey; hands and feet dull fawn. black with some infusions of brown into the dark brown region E. macrurus (Mjöberg, 1816) from the southern wet tropics in anterior to the eye. Queensland, Australia, generally south and west of Cairns, Distribution: E. rosswellingtoni sp. nov. is known from high North Queensland are readily separated from the other five altitude areas of the central cordillera of New Guinea, wholly herein recognized species by having a brownish-grey colour within the territory of Papua New Guinea, from just inside the above, with a slight rufous tinge at the anterior end only. The border with Irian Jaya (Indonesia), east through well-known high face is rufous fawn or just yellowish brown, depending on the points like Mount Wilhelm and further east to at least Mount individual, with a square black patch across each eye and Erimbari in the Eastern Highlands Province of Papua New nowhere near the ears. This patch is notable for having a well Guinea and south to at least Mt. Sisa, Southern Highlands defined black triangle jutting out from above the rear of the eye, Province, Papua New Guinea. the apex pointing towards the crown. Etymology: Named in honour of Cliff Ross Wellington, better Ears are rather long and narrow, evenly oval, naked and greyish known as Ross Wellington, of New South Wales, Australia in on the lobes and rich pink within. Back-hairs are slaty grey for recognition of a massive contribution to herpetology and wildlife five sixths of their length, their tips a brownish-grey colour. Chest conservation in Australia over some decades and who also and belly pale yellowish white, the bases of the hairs slate- publicly took a stand against taxonomic vandalism by Wolfgang coloured. Wuster and his gang of thieves, via a formal written submission to the ICZN (Wellington 2015) and other critically important Arms and legs brown on top, light (whitish) underneath; hands works for the benefit of science. and feet a light pinkish colour. E. richardwellsi sp. nov. from the northern wet tropics generally EUDROMICIA JACKYHOSERAE SP. NOV. from north of Port Douglas and south of Cape Tribulation, with a LSID urn:lsid:zoobank.org:act:81DB7D79-205A-42F3-9878- distribution centred on the Mount Lewis area is similar in most 8FA85727428D respects to E. macrurus, which it would otherwise be identified Holotype: A preserved specimen at the Yale Peabody Museum. as. E. richardwellsi sp. nov. can however be separated from E. New Haven, Connecticut, United States of America, Vertebrate macrurus by having significantly less-well defined black around Zoology Division, Mammalogy, specimen number 001701, the eye, being peppered in appearance and not boldly collected from Nassau Range, Irian Jaya, Indonesia, Latitude - demarcated from the adjacent brownish fur as seen in E. 4.20 S., Longitude 137.00 E. This facility allows access to its macrurus. The jowls region in E. richardwellsi sp. nov. is yellow holdings. rather than whitish in E. macrurus. Diagnosis: The species complex, until now regarded as The species E. adelynhoserae sp. nov. from the Milne Bay putative Eudromicia (formerly Cercartetus) caudatus (Milne- Province, Papua New Guinea is readily separated from the other Edwards, 1877) (herein restricted to the type region of the type species by having a distinctly steel-greyish colour above, locality, being the Arfak Mountains in western Papua, contrasting with white undersides and jowls. The upper surface Indonesia), including E. macrurus (Mjöberg, 1816) (from the has a somewhat peppered appearance. The ears are blackish southern wet tropics in Queensland, Australia) and the four grey, the dark colour extending almost all the way into the visible newly described species, all formerly treated as populations of ear, with inner regions a pale orangeish colour. The black E. caudatus, namely E. richardwellsi sp. nov. (from the northern patches over the eyes are elliptical in shape and extend from the wet tropics, Queensland Australia), E. adelynhoserae sp. nov. snout. There is an ill defined line of blackish grey running from (from Milne Bay, PNG), E. rosswellingtoni sp. nov. (from the the mid-snout between the eyes, further back to the crown. central ranges of New Guinea, generally near Mount Wilhelm The species E. rosswellingtoni sp. nov. from the region around and west to the Irian Jaya border), E. jackyhoserae sp. nov. Mount Wilhelm and including nearby elevated parts of Eastern (from Nassau Range, Irian Jaya) and E. doriskuenae sp. nov. Highlands and Southern Highlands provinces of Papua New from the high altitude areas of the Finisterre Range in Madang Guinea, west to the Irian Jaya border is readily separated from and Morobe Provinces, PNG, are readily separated from all the other species by having brown hair on the upper body, ears other members of the genus by having a large size as in being that are brownish grey at the outer edges and becoming orange more than 85 mm head and body length; lower leg length more further in, feet are pink and the pink of the snout almost extends than 20 mm long and hind leg more than 13 mm long. The tail is to the eyes. The belly is beige and the lower flanks have well also long and more than 120 mm in length, slightly thickened defined demarcation between darker upper parts and lighter and furred at the base (separating it from the superficially similar underparts, which do not run in an even line. Pogonomys Milne-Edwards, 1877). The dark mark through the The species E. jackyhoserae sp. nov. from the Nassau Range in eye is obvious and there are large crinkly ears. This is also a Irian Jaya (West Papua), Indonesia is similar in most respects to genus-level diagnosis for all species within Eudromicia. E. rosswellingtoni sp. nov. as described above, but can be Eudromicia caudatus (Milne-Edwards, 1877) herein restricted to separated from that species by having orange-pink feet and the type region of the type locality, being the Arfak Mountains in mottled darker patches or blotches on the sides of the face and western Papua, Indonesia and all other species in the complex, forelimbs. The ears have greyish tips, but are otherwise are easily separated from one another on colouration. yellowish in colour further in. Eudromicia caudatus (Milne-Edwards, 1877) is readily separated The belly is beige and the lower flanks have well defined from the other five herein recognized species by having a dull demarcation between darker upper parts and lighter underparts, rufous colour above. The face is rufous fawn, with two broad which do not run in an even line. The upper limbs and lower black bands passing from the nose through the eyes, but not flanks are dark brown with well defined lighter edges, forming quite reaching the ears. Ears are rather long and narrow, evenly large patches and quite distinct from the brownish grey dorsal oval, naked. Back-hairs are slaty grey for five sixths of their surfaces. length, their tips dull rufous. Chest and belly pale yellowish

Hoser 2020 - Australasian Journal of Herpetology 42:11-22. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 20 Australasian Journal of Herpetology white, the bases of the hairs slate-coloured. around the eye, the black marking is dark brown rather than Arms and legs grey; hands and feet dull fawn. black with some infusions of brown into the dark brown region E. macrurus (Mjöberg, 1816) from the southern wet tropics in anterior to the eye. Queensland, Australia, generally south and west of Cairns, Distribution: E. jackyhoserae sp. nov. is known only from the North Queensland are readily separated from the other five Nassau Range, Irian Jaya, Indonesia, but is probably found at herein recognized species by having a brownish-grey colour, other high elevation locations nearby. with a slight rufous tinge at the anterior end only. The face is Etymology: Named in honour of my eldest daughter, Jacky rufous fawn or just yellowish brown above, depending on the Hoser of Park Orchards, (Melbourne), Victoria, Australia in individual, with a square black patch across each eye and recognition of over 18 years of significant contributions to wildlife nowhere near the ears. This patch is notable for having a well conservation in Australia, including her work with Snakebusters: defined black triangle jutting out from above the rear of the eye, Australia’s best reptiles shows, being the only hands on wildlife the apex pointing towards the crown. shows in Australia that let people hold the animals. Ears are rather long and narrow, evenly oval, naked and greyish Jacky Hoser also made significant contributions to several years on the lobes and rich pink within. Back-hairs are slaty grey for research on the Mountain Pygmy Possums of the genus five sixths of their length, their tips a brownish-grey colour. Chest Burramys Broom (1896) in the snowfields of Victoria, Australia. and belly pale yellowish white, the bases of the hairs slate- EUDROMICIA DORISKUENAE SP. NOV. coloured. LSID urn:lsid:zoobank.org:act:115D1E32-E68D-48B7-88F6- Arms and legs brown on top, light (whitish) underneath; hands 8633DBA7F01B and feet a light pinkish colour. Holotype: A preserved specimen at the Australian Museum, E. richardwellsi sp. nov. from the northern wet tropics generally Sydney, New South Wales, Australia, specimen number from north of Port Douglas and south of Cape Tribulation, with a M.21654, collected at Nokopo, Madang Province, Papua New distribution centred on the Mount Lewis area is similar in most Guinea, Latitude -5.95 S., Longitude 146.60 E. This facility respects to E. macrurus, which it would otherwise be identified allows access to its holdings. as. E. richardwellsi sp. nov. can however be separated from E. Diagnosis: The species complex, until now regarded as macrurus by having significantly less-well defined black around putative Eudromicia (formerly Cercartetus) caudatus (Milne- the eye, being peppered in appearance and not boldly Edwards, 1877) (herein restricted to the type region of the type demarcated from the adjacent brownish fur as seen in E. locality, being the Arfak Mountains in western Papua, macrurus. The jowls region in E. richardwellsi sp. nov. is yellow Indonesia), including E. macrurus (Mjöberg, 1816) (from the rather than whitish in E. macrurus. southern wet tropics in Queensland, Australia) and the four The species E. adelynhoserae sp. nov. from the Milne Bay newly described species, all formerly treated as populations of Province, Papua New Guinea is readily separated from the other E. caudatus, namely E. richardwellsi sp. nov. (from the northern species by having a distinctly steel-greyish colour above, wet tropics, Queensland Australia), E. adelynhoserae sp. nov. contrasting with white undersides and jowls. The upper surface (from Milne Bay, PNG), E. rosswellingtoni sp. nov. (from the has a somewhat peppered appearance. The ears are blackish central ranges of New Guinea, generally near Mount Wilhelm grey, the dark colour extending almost all the way into the visible and west to the Irian Jaya border), E. jackyhoserae sp. nov. ear, with inner regions a pale orangeish colour. The black (from Nassau Range, Irian Jaya) and E. doriskuenae sp. nov. patches over the eyes are elliptical in shape and extend from the from the high altitude areas of the Finisterre Range in Madang snout. There is an ill defined line of blackish grey running from and Morobe Provinces, PNG, are readily separated from all the mid-snout between the eyes, further back to the crown. other members of the genus by having a large size as in being The species E. rosswellingtoni sp. nov. from the region around more than 85 mm head and body length; lower leg length more Mount Wilhelm and including nearby elevated parts of Eastern than 20 mm long and hind leg more than 13 mm long. The tail is Highlands and Southern Highlands provinces of Papua New also long and more than 120 mm in length, slightly thickened Guinea, west to the Irian Jaya border is readily separated from and furred at the base (separating it from the superficially similar the other species by having brown hair on the upper body, ears Pogonomys Milne-Edwards, 1877). The dark mark through the that are brownish grey at the outer edges and becoming orange eye is obvious and there are large crinkly ears. This is also a further in, feet are pink and the pink of the snout almost extends genus-level diagnosis for all species within Eudromicia. to the eyes. The belly is beige and the lower flanks have well Eudromicia caudatus (Milne-Edwards, 1877) herein restricted to defined demarcation between darker upper parts and lighter the type region of the type locality, being the Arfak Mountains in underparts, which do not run in an even line. western Papua, Indonesia and all other species in the complex, The species E. jackyhoserae sp. nov. from the Nassau Range in are easily separated from one another on colouration. Irian Jaya (West Papua), Indonesia is similar in most respects to Eudromicia caudatus (Milne-Edwards, 1877) is readily separated E. rosswellingtoni sp. nov. as described above, but can be from the other five herein recognized species by having a dull separated from that species by having orange-pink feet and rufous colour above. The face is rufous fawn, with two broad mottled darker patches or blotches on the sides of the face and black bands passing from the nose through the eyes, but not forelimbs. The ears have greyish tips, but are otherwise quite reaching the ears. Ears are rather long and narrow, evenly yellowish in colour further in. oval, naked. Back-hairs are slaty grey for five sixths of their The belly is beige and the lower flanks have well defined length, their tips dull rufous. Chest and belly pale yellowish demarcation between darker upper parts and lighter underparts, white, the bases of the hairs slate-coloured. which do not run in an even line. The upper limbs and lower Arms and legs grey; hands and feet dull fawn. flanks are dark brown with well defined lighter edges, forming E. macrurus (Mjöberg, 1816) from the southern wet tropics in large patches and quite distinct from the brownish grey dorsal Queensland, Australia, generally south and west of Cairns, surfaces. North Queensland are readily separated from the other five The species E. doriskuenae sp. nov. from the high altitude areas herein recognized species by having a brownish-grey colour of the Finisterre Range in Madang and Morobe Provinces, PNG above, with a slight rufous tinge at the anterior end only. The is similar in most respects to E. rosswellingtoni sp. nov. however face is rufous fawn or just yellowish brown, depending on the can be differentiated from that species and in turn all others, by individual, with a square black patch across each eye and the fact that demarcation between darker upper parts and lighter nowhere near the ears. This patch is notable for having a well underparts is not well defined; inner ears are orange in colour defined black triangle jutting out from above the rear of the eye, except for the very outer tips which have grey outer edges; the apex pointing towards the crown.

Ears are rather long and narrow, evenly oval, naked and greyish REFERENCES CITED on the lobes and rich pink within. Back-hairs are slaty grey for Anonymous 2007. Two new mammals found in Indonesian ‘lost five sixths of their length, their tips a brownish-grey colour. Chest world’: green group. Media release dated 17 December, posted and belly pale yellowish white, the bases of the hairs slate- online at: coloured. Arms and legs brown on top, light (whitish) https://26sep.net/news_details.php?lng=english&sid=36002 underneath; hands and feet a light pinkish colour. Aplin, K., Dickman, C., Salas, L., Burnett, S. and Winter, J. E. richardwellsi sp. nov. from the northern wet tropics generally 2016. Cercartetus caudatus. IUCN Red List of Threatened from north of Port Douglas and south of Cape Tribulation, with a Species (version 3.1). International Union for Conservation of distribution centred on the Mount Lewis area is similar in most Nature and Natural Resources, Online. respects to E. macrurus, which it would otherwise be identified Bannister, J. L., Calaby, J. H., Dawson, L. J., Ling, J. K., as. E. richardwellsi sp. nov. can however be separated from E. Mahoney, J. A., McKay, G. M., Richardson, B. J., Ride, W. D. L. macrurus by having significantly less-well defined black around and Walton, D. M. 1988. Zoological Catalogue of Australia, the eye, being peppered in appearance and not boldly Volume 5, Mammals. Australian Government Publishing Service, demarcated from the adjacent brownish fur as seen in E. Canberra, ACT, Australia:X+274 pp. macrurus. The jowls region in E. richardwellsi sp. nov. is yellow rather than whitish in E. macrurus. Bell, T. 1828. Description of a new species of Phalangista. Trans The species E. adelynhoserae sp. nov. from the Milne Bay Linn. Soc. London 16:121-127. Province, Papua New Guinea is readily separated from the other Brammell, J. and Archer, M. 1997. A new Oligocene-Miocene species by having a distinctly steel-greyish colour above, species of Burramys (Marsupialia, Burramyidae) from contrasting with white undersides and jowls. The upper surface Riversleigh, northwestern Queensland. Memoirs of the has a somewhat peppered appearance. The ears are blackish Queensland Museum 41(2):247-268. grey, the dark colour extending almost all the way into the visible Broom, R. 1895. On a new fossil mammal allied to ear, with inner regions a pale orangeish colour. The black Hypsiprymnus but resembling in some points the patches over the eyes are elliptical in shape and extend from the Plagiaulacidae. Abstr. Proc.Linn.Soc. N.S.W. (1 June):26:11. snout. There is an ill defined line of blackish grey running from Broom, R. 1896a. On a small fossil marsupial with large grooved the mid-snout between the eyes, further back to the crown. premolars. Proc. of the Linnaean Soc. N.S.W., 2(10):563-567. The species E. rosswellingtoni sp. nov. from the region around Broom, R. 1896b. On a small fossil marsupial allied to Petaurus. Mount Wilhelm and including nearby elevated parts of Eastern Proceedings of the Linnaean Soc. N.S.W., 2(10):568-570. Highlands and Southern Highlands provinces of Papua New Brongersma, L. D. 1953. Notes on New Guinean reptiles and Guinea, west to the Irian Jaya border is readily separated from amphibians. II. Proceedings Koninklijke Nederlandsche the other species by having brown hair on the upper body, ears Akademie van Wetenschappen, Amsterdam (C) 56:317-325. that are brownish grey at the outer edges and becoming orange further in, feet are pink and the pink of the snout almost extends Collins, L. R. 1973. Monotremes and Marsupials. Smithsonian to the eyes. The belly is beige and the lower flanks have well Institution Press. City of Washington, DC, USA. defined demarcation between darker upper parts and lighter Bryant, L. M. and Krosch, M. N. 2016. Lines in the land: a review underparts, which do not run in an even line. of evidence for eastern Australia’s major biogeographical The species E. jackyhoserae sp. nov. from the Nassau Range in barriers to closed forest taxa. Biological Journal of the Linnean Irian Jaya (West Papua), Indonesia is similar in most respects to Society, 2016:27 pp. (online) E. rosswellingtoni sp. nov. as described above, but can be Desmarest, A. G. 1818. Nouveau Dictionairie d’Histoire separated from that species by having orange-pink feet and Naturelle. Deterville, Tome 25:477. mottled darker patches or blotches on the sides of the face and Desmarest, A. G. 1820. Mammalogie ou description des forelimbs. The ears have greyish tips, but are otherwise especes de mammiferes. Encyclopedic Methodique Histoire yellowish in colour further in. Naturelle. Volume I, (Mme Veuve Agasse: Paris). 1:268. The belly is beige and the lower flanks have well defined Dwyer, P. D. 1977. Notes on Antechinus and Cercartetus demarcation between darker upper parts and lighter underparts, (Marsupialia) in the New Guinea Highlands. Proceedings of the which do not run in an even line. The upper limbs and lower Royal Society of Queensland 88:69-73. flanks are dark brown with well defined lighter edges, forming Eldridge, M. D. B., Potter, S., Helgen, K. M., Sinaga, M. H., large patches and quite distinct from the brownish grey dorsal Aplin, K. P., Flannery, T. F. and Johnson, R. N. 2018. surfaces. Phylogenetic analysis of the tree-kangaroos (Dendrolagus) The species E. doriskuenae sp. nov. from the high altitude areas reveals multiple divergent lineages within New Guinea. of the Finisterre Range in Madang and Morobe Provinces, PNG Molecular Phylogenetics and Evolution, 127:588-599 (online). is similar in most respects to E. rosswellingtoni sp. nov. however Flannery, T. F. 1995. The Mammals of New Guinea, 2nd edition. can be differentiated from that species and in turn all others, by Reed Books, Sydney, Australia. the fact that demarcation between darker upper parts and lighter Gloger, C. W. L. 1841. Gemeinnütziges Hand- und Hilfsbuch der underparts is not well defined; inner ears are orange in colour Naturgeschichte. Für gebildete Leser aller Stände, besonders except for the very outer tips which have grey outer edges; für die reifere Jugend und ihre Lehrer. Erster Band, enthaltend around the eye, the black marking is dark brown rather than die erste Hälfte der Naturgeschichte der Thiere, nebst black with some infusions of brown into the dark brown region erfahrungsmässigen Andeutungen über den gegenwärtigen anterior to the eye. Zustand und Erfolg des Unterrichts in dieser Wissenschaft, Distribution: E. doriskuenae sp. nov. is only known from the namentlich auf Gymnasien, und Vorrschlägen über fernere high altitude areas of the Finisterre Range in Madang and Einrichtung desselben im Verhaltnisse zu seinem Wirklichen Morobe Provinces, Papua New Guinea in a region generally Zwecke. A. Schulz & Comp., Breslau. Vol. Hefte 1-5:400 pp. bounded by the Markham and Ramu River basins to the south Gould, J. 1845. Untitled abstract. Proceedings of the Zoological and the coast to the north. Society of London 1845:2. Etymology: Named in honour of Doris Kuen of Donvale, Gray, J. E. 1841. Description of some new species and four new Victoria, Australia, for services to conservation, through her genera of reptiles from Western Australia, discovered by John vitally important work in maintaining the structure, electrical Gould, Esq. Ann. Mag. Nat. Hist. (1)7:86-91. fittings, plumbing, repairs to motor vehicles at urgent times and other important tasks at the Snakebusters, wildlife displays and Groves, C. P., Wilson, D. E. and Reeder, D. M. (eds.) 2005. scientific research facility in Park Orchards, Victoria, Australia. Mammal Species of the World: A Taxonomic and Geographic Hoser 2020 - Australasian Journal of Herpetology 42:11-22. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 22 Australasian Journal of Herpetology

Reference (3rd ed.). Johns Hopkins Uni Press, Baltimore, USA. over de natuurlijke geschiedenis der Nederlandsche Harris, J. M. 2006. The discovery and early natural history of the overzeesche bezittingen, door leden der Natuurkundige eastern pygmy-possum, Cercartetus nanus (Geoffroy and Commissie in Indië en andere schrijvers. Luchtmans, Leiden:1- Desmarest, 1817). Proceedings of the Linnean Society of New 57. South Wales. 127:107-124. Osborne, M. J. and Christidis, L. 2002. Systematics and Harris, J. M. 2010. The natural history, conservation status and biogeography of pygmy possums (Burramyidae: Cercartetus). ecology of the eastern pygmy-possum (Cercartetus nanus). PhD Journal of Australian Zoology 50(1):25-37. thesis, Southern Cross University, Lismore, NSW. Online at: Osborne, M. J., Norman, J. A., Christidis, L. and Murray, N. D. https://epubs.scu.edu.au/cgi/ 2000. Genetic distinctiveness of isolated populations of an viewcontent.cgi?article=1294&context=theses endangered marsupial, the mountain pygmy-possum, Burramys Hoser, R. T. 1989. Australian Reptiles and Frogs. Pierson parvus. Molecular Ecology 9:609-613. Publishing, Mosman, NSW, Australia:238 pp. Pestell, A. J. L., Cooper, S. J. B., Saint, K. and Petit, S. 2008. Hoser, R. T. 1991. Endangered Animals of Australia. Pierson Genetic structure of the western pygmy possum Cercartetus Publishing, Mosman, NSW, Australia:240 pp. concinnus Gould (Marsupialia: Burramyidae) based on Hoser, R. T. 2015. Australian agamids: Eighteen new species mitochondrial DNA. Australian Mammalogy. 29(2):191-200. from the genera Amphibolurus Wagler, 1830, Lophognathus Parker, H. W. 1940. The Australian frogs of the family Gray, 1842, Rankinia Wells and Wellington, 1984, Diporiphora Leptodactylidae. Novit. Zool. 42:1-106. Gray, 1842, Tympanocryptis Peters, 1863, as well as three new Peters, W. C. H. 1863. 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A classification of the expeditions to Australia 1910-13. 2 Mammals from Queensland Amphibia and Reptilia of Australia. Australian Journal of 2. On a new genus and species of Marsupial. K. svenska Herpetology Supplementary Series 1:1-61. VetenskAkad. Handl., 52(2):13-20. Zaczek, Z. 2019. Former PM Kevin Rudd says Australia should Moritz, C., Joseph, L. and Adams, M. 1993. Cryptic diversity in drastically increase its population to 50 MILLION to make sure nation can defend itself in the face of threats from China. Daily an endemic rainforest skink (Gnypetoscincus queenslandiae). Hoser 2020 - Australasian Journal of Herpetology 42:11-22. Biodiversity and Conservation 2:412-425. Mail (Australia) (27 November), published online at: https:// Available online atwww.dailymail.co.uk/news/article-7729689/Kevin-Rudd-says- www.herp.net Müller, S. 1840. Over de zoogdieren van den Indischen Archipel. Australia-increase-population-50-MILLION-defend-China.html In: Temminck, C.J., van der Hoek, J. (Editors), Verhandelingen Copyright- Kotabi PublishingCONFLICTS - All rights OF INTEREST reserved - NONE. Australasian Journal of Herpetology 23 Australasian Journal of Herpetology 42:23-30. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

A new subgenus, three new species and one new subspecies of Ring- tailed Possums (Marsupialia: Petauridae) from the north of Australia.

LSIDURN:LSID:ZOOBANK.ORG:PUB:3C179FF3-EF6C-4D64-93F8-12D1CFB714B0

RAYMOND T. HOSER

LSID urn:lsid:zoobank.org:author:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 8 April 2020, Accepted 14 April 2020, Published 25 April 2020.

ABSTRACT A review of the taxonomy of the Australian Ring-tailed Possums in the associated putative genera Petropseudes Thomas, 1923 (Rock Ringtail Possums) of Australia’s dry tropics region and Pseudochirops Matschie, 1915, of Australia’s wet tropics region was conducted. Including just two putative species Petropseudes dahli (Collett, 1895) and Pseudochirops archeri (Collett, 1884) (Green Ringtail Possum), the study involved inspection of specimens of each form across their known distributions. In the case of both putative taxa, there was found to be significant variation in morphology in regionally disjunct populations located across well-known biogeographical barriers of known antiquity. As a result, Petropseudes dahli (Collett, 1895) was found to consist of no less than four species. Three are formally named for the first time in accordance with the International Code of Zoological Nomenclature (Ride et al. 1999). Meredith et al. (2010) noted a six million year divergence of this species group from nominate Pseudochirops. On this basis Petropseudes is designated a subgenus within Pseudochirops as originally intended by Thomas (1923). Pseudochirops archeri (Collett, 1884) of North Queensland was found to include two morphologically divergent and reproductively isolated populations in close proximity, but on either side of the so-called Black Mountain Corridor (or barrier) as identified by Bryant and Krosch (2016). The unnamed population is herein formally named as a new subspecies. The Australian endemic, Pseudochirops archeri is not only divergent from the other known members of the genus Pseudochirops Matschie, 1915, (all from the New Guinea subregion), but also the taxon currently called Petropseudes dahli herein treated as being within Pseudochirops. Meredith et al. (2010) estimated a divergence of Pseudochirops archeri from nearest common ancestors of the other species at about eight MYA and so a new subgenus is erected to accommodate this species. Keywords: Mammals; Marsupial; Petauridae; taxonomy; nomenclature; possum; Ring-tailed possum; Petropseudes; Pseudochirops; Pseudocheirus; dahli; archeri; Australia; Queensland; Northern Territory; Western Australia; New Guinea; wet tropics; dry tropics; subgenus; new subgenus; Sloppossum; new species; fiacummingae; jamesbondi; waddamaddawidyu; new subspecies; chrismaxwelli.

INTRODUCTION the same putative species at locations more than 1,000 km Over a period of more than 30 years doing fieldwork involving apart. Hence it came as a surprise that one of the two wildlife across northern parts of Australia, specimens of Rock populations was effectively unrecognized taxonomically, even Ringtail Possums Petropseudes dahli (Collett, 1895) were though the two relevant populations were split across a well- observed in the Northern Territory and Western Australia both in known biogeographical barrier near the Northern Territory / the wild and in captivity. Western Australian border. It was observed by myself and without fanfare that they differed In fact the nominate form, originally described as Pseudochirops significantly in morphology, as one would expect when seeing dahli Collett, 1895 with a type locality of Mary River, Northern Hoser 2020 - Australasian Journal of Herpetology 42:23-30. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 24 Australasian Journal of Herpetology

Territory (Kakadu area) is the only available name for any genus, necessitating the elevation of the new subgenus populations of this putative species (Bannister et al. 1988). Sloppossum subgen. nov. to full genus status. Hence it was decided to undertake a review of the putative I note that Sloppossum subgen. nov. is in fact more divergent species across the known range from north-west Queensland, than Petropseudes. across the Northern Territory and into the Kimberley division of MATERIALS, METHODS AND RESULTS north-west Western Australia, to ascertain the status of given Before a decision is made to name any new taxon, reasonable populations and whether any should be accorded species or steps must be taken to ensure that it is justified on all relevant subspecies-level recognition. grounds, including that it is morphologically, genetically and Counter to this proposition is whether or not clines exist between reproductively isolated from their nearest relative and to a regional forms, or did so in the recent past and in the process of sufficient degree to be of taxonomic significance. investigation it soon became apparent that none existed. A further relevant question to ask is should the reproductively With each of four main known populations being morphologically isolated and morphologically divergent entities be labelled as divergent and having been reproductively isolated for some subspecies, full species, or potentially higher level again. geological time, it then became a matter of deciding at what Key literature relevant to the taxonomic and nomenclatural level of classification each should be identified as. conclusions within this paper include While such a decision is subjective, due to the likelihood that divergences between populations are likely to be measured in Bannister et al. (1988), Bryant and Krosch (2016), Collett (1884), the millions of years, I had no hesitation in deciding to name the Collins (1973), Eldridge et al. (2018), Flannery (1995), three main unnamed populations as species. Goodfellow (1993), Groves et al. (2005), Iredale and Troughton (1934), Jones et al. (2006), Laurance (1990), Lawlor (1979), Obviously this decision was also made after consulting all Matschie (1915), Maxwell et al. (1986), Menkhorst (2001), relevant literature in terms of this and similar species and other Meredith et al. (2010), Menzies (1991), Moritz et al. (1993), vertebrate species in the same regions and affected by the Potter et al. (2014), Ride (1970), Ride et al. (1999), Runcie same biogeographical features and timelines (see for example (1999), Strahan (as editor) (1988), Thomas (1888, 1922, 1923), Potter et al. 2014). van Ufford and Cloos (2005), Vaughan (1986) and sources cited In the case of the three putative candidate taxa, the well-known therein (duplicitous references not necessarily included). biogeographical barriers had known antiquity. Live and dead specimens as well as available bone specimens, As a result, Petropseudes dahli (Collett, 1895) was found to were examined as was other relevant material, including past consist of four species. Three are formally named for the first climate data for the relevant regions, sea level depths, and other time in accordance with the International Code of Zoological important information. Nomenclature (Ride et al. 1999). In summary, as inferred already, the genetic, geological, Meredith et al. (2010) noted a six million year divergence of this historical and morphological evidence clearly showed that both species group from nominate Pseudochirops which is a relevant of Petropseudes dahli (Collett, 1895) (Rock Ringtail Possums) fact that would be hard to miss in any review of the putative and Pseudochirops archeri (Collett, 1884) (Green Ringtail species. Possum) as currently understood required species-level On this basis Petropseudes Thomas, 1923 is designated a divisions and a refinement of their genus-level placements. subgenus within Pseudochirops. SUMMARY OF THE RESULTS I note that this is exactly as originally intended by Thomas Petropseudes dahli (Collett, 1895) (Rock Ringtail Possums), are (1923) who in fact created the name Petropseudes. found across the dry tropics of Australia from north-west Another species I had the good fortune to observe in the wild on Queensland in the vicinity of Lawn Hill, along the southern shore a number of occasions was Pseudochirops archeri (Collett, of the Gulf of Carpentaria and across the top end of the Northern 1884) of North Queensland, better-known as the Green Ringtail Territory to the western Kimberley Ranges in Western Australia. Possum. However a closer inspection of collection records and reliable It was also reviewed and found to include two morphologically sightings shows that the putative species is in fact confined to divergent and reproductively isolated populations in close rocky regions, including outliers and nowhere else, in effect proximity, but on either side of the so-called Black Mountain confining the distribution to four main areas in drier parts of Corridor (or barrier) as identified by Bryant and Krosch (2016). northern Australia. With the type locality for the species being the Herbert River These are: area of north Queensland, within the southern wet tropics area, this meant that the northern population from the Mount Lewis 1/ The type locality around the Arnhem Land escarpment in the area was effectively unnamed. middle of the top end of the Northern Territory. Notwithstanding the known antiquity of the P. archeri lineage as 2/ Rocky parts of far north-west Queensland, extending along found by Meredith et al. (2010), the morphological divergence of the western shoreline of the Gulf of Carpentaria, into the the two populations and geographical proximity of them has led Northern Territory. me to take a conservative position with regards to taxonomy and 3/ Groote Eylandt. the unnamed form is herein named as a new subspecies. 3/ The Western part of the Kimberley division of Western The Australian endemic, Pseudochirops archeri is not only Australia. divergent from the other known members of the genus The two westernmost populations are well separated by the Pseudochirops Matschie, 1915, (all from the New Guinea corridor of the Victoria River system in the north-west of the subregion), but also the taxon currently called Petropseudes Northern Territory and the East Kimberley. dahli herein treated as being within Pseudochirops. There are some alleged sighting records within the eastern part Meredith et al. (2010) estimated a divergence of Pseudochirops of this zone, but even allowing them to be accurate and reliable, archeri from nearest common ancestors of the other species at (which is a leap of faith in itself, noting there are other about eight MYA and so a new subgenus is erected to morphologically similar species in the same region) one still accommodate this species. finds a zone of several hundred kilometres in a straight line that Based on the published divergences within Meredith et al. has never had any putative P. dahli recorded in any way, even (2010), if one were to continue to regard Petropseudes as a though it includes areas heavily collected by State Museums. valid genus, then one would have no choice but to treat the Hence the absence of specimen records must be attributed to species Pseudochirops archeri as being within a separate an absence of specimens as opposed to an absence of

The gap between the central (NT) and mainland eastern NT/ Pseudochirops archeri (Collett, 1884) of North Queensland was Queensland populations corresponds to a flat zone of unsuitable also scrutinized as it was known to have two populations on habitat stretching at least 100 km (being east of the Arnhem either side of the so-called Black Mountain Corridor (or barrier) Land escarpment), this area not expecting to have populations as identified by Bryant and Krosch (2016), being a barrier I have and for which no sighting or other records exist. exploited to find and identify a series of new and previously Hence, even in the absence of inspection of specimens, there is cryptic species as seen for example in Hoser (2016a, 2016b) or an inescapable conclusion that the three main mainland see Moritz et al. (1993). populations must be reproductively separated. Like Pseudochirops (Petropseudes) dahli (Collett, 1895), P. There is no evidence to suggest that actions by humans since archeri exhibits strong habitat fidelity and there is no evidence to first settlement 40K MYA to present or changes wrought by ice suggest it can breach the Black Mountain Corridor in the current age climate fluctuations have altered the biogeography of the interglacial, let alone in glacial periods when the same corridor relevant part of Australia to change the current status quo with would be wider and drier and less able to be breached by a regards to population mobility. rainforest obligate species. I also note that there is no evidence this putative species (P. Jones et al. (2006) found the that in terms of diet for the putative dahli) has ever made it to New Guinea, even though its current species P. archeri that “Over 50% of tree use was from only four distribution sits proximal to a historical land bridge to southern tree species, Aleurites rockinghamensis, Ficus fraseri, Arytera New Guinea. divaricata and Ficus copiosa.”, all of which are rainforest This is further evidence supporting the contention of habitat obligate species not found in the region of the Black Mountain fidelity for the putative species and that dispersal is not likely Gap (as per ALA records of all types), or unlikely to have been between known and well-spaced populations. there in a glacial phase, when the relevant area would have By straight line measurement the Groote Eylandt population is been even drier than present. about 100 km from the nearest known population on the Inspection of specimens of P. archeri found P. archeri to mainland of Australia, situated to the south-west. include two morphologically divergent and reproductively Groote Eylandt is known to have endemic species and isolated populations in close proximity, but on either side of the subspecies of reptile and mammals for which divergences are so-called Black Mountain Corridor (or barrier) as identified by well-known and with this in mind specimens of P. dahli from Bryant and Krosch (2016). The unnamed population is herein there were inspected, with a view to potentially identifying a formally named as a new subspecies P. archeri chrismaxwelli fourth species. subsp. nov., being within the subgenus Sloppossum subgen. Although they were morphologically most similar to the nov. as explained below. specimens from the Gulf of Carpentaria, including in terms of The Australian endemic, Pseudochirops archeri is not only the diagnostic features for that species, they are easily divergent from the other known members of the genus separable and have therefore been named herein as a separate Pseudochirops Matschie, 1915, (all from the New Guinea and newly named species. subregion), but also the taxon currently called Petropseudes I also note that biogeographically, the fauna of Groote Eylandt is dahli herein treated as being within Pseudochirops due to its more closely affiliated with that of the Arnhem Land escarpment affinity to that genus to the exclusion of all others. than it is to that from the southern Gulf of Carpentaria, which Meredith et al. (2010) estimated a divergence of Pseudochirops implies further morphological divergence of the Groote Eylandt archeri from nearest common ancestors of the other species at form as compared to that from the Arnhem Land region. about eight MYA. Significantly the taxonomic division of the original putative The species is also significantly morphologically divergent to all species Petropseudes dahli is across exactly the same others within Pseudochirops and so a new subgenus is erected biogeographical barriers as used to divide the putative species to accommodate this species. Odatria (Kimberleyvaranus) glebopalma (Mitchell, 1955) by The subgenus Sloppossum subgen. nov. is formally named and Hoser (2018) who in turn referred to Hoser (2014) defined in this paper before the species and subspecies (in that As already mentioned, Meredith et al. (2010) noted a six million order) in accordance with online requests and directives by the year divergence of this species group from nominate ICZN and managers of their name repository, “Zoobank”. Pseudochirops which is a relevant fact that would be hard to INFORMATION RELEVANT TO THE FORMAL DESCRIPTIONS miss in any review of the species. THAT FOLLOW On this basis Petropseudes is herein designated a subgenus There is no conflict of interest in terms of this paper or the within Pseudochirops. conclusions arrived at herein. I note that this is exactly as originally intended by Thomas Several people including anonymous peer reviewers who (1923) who in fact created the name Petropseudes. revised the manuscript prior to publication are also thanked as In effect this means that the Rock Ringtail Possums of the dry are relevant staff at museums who made specimens and tropics of northern Australia, now consist of four species, three records available in line with international obligations. of which are named for the first time in accordance with the In terms of the following formal descriptions, spellings should International Code of Zoological Nomenclature (Ride et al. not be altered in any way for any purpose unless expressly and 1999). exclusively called for by the rules governing Zoological These are: Nomenclature as administered by the International Commission 1/ Pseudochirops (Petropseudes) dahli (Collett, 1895) herein of Zoological Nomenclature. restricted to the Arnhem Land escarpment and immediately In the unlikely event two or more newly named taxa are deemed adjacent outliers in the Northern Territory. conspecific by a first reviser, then the name to be used and 2/ Pseudochirops (Petropseudes) fiacummingae sp. nov. from retained is that which first appears in this paper by way of page the west and north of the Kimberley Division of Western priority and as listed in the abstract keywords. Australia. Some material in descriptions for taxa may be repeated for other 3. Pseudochirops (Petropseudes) jamesbondi sp. nov. from taxa in this paper and this is necessary to ensure each fully north-west Queensland and nearby parts of far eastern Northern complies with the provisions of the International Code of Territory, generally south and west of the Gulf of Carpentaria. Zoological Nomenclature (Fourth edition) (Ride et al. 1999) as 4. Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. amended online since. from Groote Eylandt, Northern Territory. Material downloaded from the internet and cited anywhere in this

Hoser 2020 - Australasian Journal of Herpetology 42:23-30. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 26 Australasian Journal of Herpetology paper as being sourced online was downloaded and checked Australian Museum, Perth, Western Australia, Australia, most recently as of 10 March 2020 (including if also viewed specimen number M15934 collected from an outlier on the prior), unless otherwise stated and was accurate in terms of the southern edge of the King Leopold Ranges, Western Australia, content (as described) cited herein as of that date. Australia, Latitude -17.2 S., Longitude 124.8 E. This Unless otherwise stated explicitly, colour and other descriptions government-owned facility allows access to its holdings. apply to living adult specimens of generally good health and not Paratype: A preserved male specimen in the Western under any form of stress by means such as excessive cool, Australian Museum, Perth, Western Australia, Australia, heat, dehydration or abnormal skin reaction to chemical or other specimen number M18158 collected from an outlier on the input. southern edge of the King Leopold Ranges, Western Australia, Colour descriptions of species refer to fur colour and not skin. Australia, Latitude -17.3 S., Longitude 124.7 E While numerous texts and references were consulted prior to Diagnosis: Until now, all populations of so-called Rock Ringtail publication of this paper, the criteria used to separate the Possum have been assigned to the same putative species, relevant species or subspecies has already been spelt out and/ namely Pseudochirops (Petropseudes) dahli (Collett, 1895) with or is done so within each formal description and does not rely on a type locality of Mary River. Northern Territory, Australia. material within publications not explicitly cited herein. That form is now herein restricted to the type locality and SLOPPOSSUM SUBGEN. NOV. adjacent hills and outliers, essentially including the Arnhem LSID urn:lsid:zoobank.org:act:536D6D52-E877-4084-8D7C- Land escarpment and adjacent hills and ranges to the south and 4DA758870BE6 west. Type species: Phalangista (Pseudochirops) archeri Collett, This species and the three newly described forms herein can be 1884. readily separated from one another by consistent differences in colour of pelage on various parts of the head and body. Diagnosis: The subgenus Sloppossum subgen. nov. within the Nominate P. dahli is readily separated from the other three genus Pseudochirops Matschie, 1915 is readily separated from species by having dark grey pelage and with mid line, from all other species in the genus Pseudochirops and all other snout being distinct, thick, blackish in colour and extends past morphologically similar Petauridae by the following unique suite the crown of the head. This line becomes wider and darker of characters: Ears very short, broader than long. Tail is white- between eyes and snout and again before a line between the tipped. Body form is thick and stout. Colour greenish yellow to ears. There is a strongly dark reddish brown overlay on fur of the yellow with a slight greenish tinge, with three vague darker and upper legs. Yellowish white under forebody. The white above and two lighter dorsal stripes. Molars 1-3 are 13.6 mm. Dental series below the eye, giving the appearance of a tight circle around the nearly or quite continuous; incisor 2 elongated horizontally, its eye is very well defined. Inside of ears is salmon colour with antero-posterior diameter twice that of incisor 3. Upper edge of significant blackening towards the outer edges. lower incisor 1 is concave. The skull and dentition are superficially similar to that of the New Guinean species Pseudochirops (Petropseudes) fiacummingae sp. nov. from the Pseudochirops albertisi (Peters, 1874), but are readily separated Kimberley district of north-west Western Australia has a light from that taxon by having large and stout teeth, relative to the grey pelage with the mid line from snout distinct, thin, blackish in more typically sized dentition in the New Guinean species. colour and it goes past the crown of the head, without thickening The subgenus Sloppossum subgen. nov. can be further between the eyes and snout or again before the line between diagnosed as follows: Fur soft, close, and thick. General colour the ears. White under forebody. grizzled yellowish to greenish-yellow. Face grey, passing into The white above and below the eye giving appearance of tight greenish yellow on the crown. Eye with distinct or semi-distinct circle around eye is not well defined. Inside of ears is orange to crescentic pale yellow or white spots above and below it. Ears salmon colour with no blackening on the outer edges. Very weak very short, rounded, hairy, their posterior edges and a large spot reddish brown overlay on fur of upper legs. beneath their bases prominently white. Nape with a distinct Pseudochirops (Petropseudes) jamesbondi sp. nov. from the black central line passing down it on to the back. Back greenish southern and western edges of the Gulf of Carpentaria in yellow to yellow, with two whitish lines margining the black Queensland and the Northern Territory, Australia has a has light central one, and these again with two indistinct darker ones grey to mouse brown pelage and with the mid-line from the outside them. Chin is greyish white, chest and belly pure white snout not going past the crown, being very indistinct and is grey or off-white, the line of demarcation either sharply marked or ill- in colour, breaking up as it moves up the snout. White under defined. Limbs like back, but rather greyer. Tail is thick basally, forebody. rapidly tapering; its basal two thirds coloured like the back, its tip The white above and below the eye giving an appearance of a white. Naked part beneath tip is less than half the length of the tight circle around the eye is well defined. Inside of ears is tail. yellow-orange becoming brownish on the outer edges. Distribution: Restricted to the wet tropics bioregion of Australia There is no obvious reddish brown or orange overlay on the fur from Mount Spec, north of Townsville, north Queensland in the of upper legs save for the same slight and very feint orange south, along the coastal rainforests, generally at high elevation overlay as seen over all parts of the body. running north to the Mount Windsor Tableland in the north. Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. from Etymology: The first part of the genus name “Slop” is in Groote Eylandt, Northern Territory, Australia, is similar in most recognition of the author’s Great Dane dog, named “Slop” for respects to P. jamesbondi sp. nov. except for the obvious and services in guarding the research facility and wildlife for many strong light orange colour of the rear of the rump and the back years and the second part of the genus name is a direct usage legs (versus either not so, or only faint orange tinge in Gulf form, of the word “Possum” as the type of animal is known and hence being in line with the rest of the dorsal surface). The mid-line “Sloppossum”. marking from the snout does not go beyond the crown as in it Content: Pseudochirops (Sloppossum) archeri (Collett, 1884) terminates well before it and it is grey in colour. including both subspecies. This marking is noticeably widest at the midpoint between the PSEUDOCHIROPS (PETROPSEUDES) FIACUMMINGAE SP. eyes, appearing like an elongated diamond shape. Fur on the NOV. lower forelimbs also has a strong orange tinge, but this is only LSID urn:lsid:zoobank.org:act:B911B1A2-8A5E-4758-81A1- on the tips, rather than all over as seen on the hind limbs. FD2B5EDDFFDC All four species of possum in the subgenus Petropseudes Holotype: A preserved adult female specimen in the Western Thomas, 1923 (see above) are readily separated from all other

possums occurring in the same region in western Queensland, This species and the three newly described forms herein can be the Northern Territory and Western Australia by a combination of readily separated from one another by consistent differences in having a much shorter tail, being less than two thirds the length colour of pelage on various parts of the head and body. of the head and body as well as having very dense fur. Nominate P. dahli is readily separated from the other three Photos of Pseudochirops dahli (Collett, 1895) in life can be seen species by having dark grey pelage and with mid line, from in Strahan (1988) at page 132 (top) and online at: snout being distinct, thick, blackish in colour and extends past https://www.flickr.com/photos/88708273@N03/40093596073/ the crown of the head. This line becomes wider and darker and at: between eyes and snout and again before a line between the ears. There is a strongly dark reddish brown overlay on fur of the https://www.flickr.com/photos/128365570@N04/42868251355/ upper legs. Yellowish white under forebody. The white above and both from Arnhem Land, Northern Territory, Australia. below the eye, giving the appearance of a tight circle around the Photos of Pseudochirops (Petropseudes) fiacummingae sp. nov. eye is very well defined. Inside of ears is salmon colour with in life can be seen online at: significant blackening towards the outer edges. https://www.flickr.com/photos/mattsummerville/48131308123/ Pseudochirops (Petropseudes) jamesbondi sp. nov. from the from Prince Regent, Western Australia, Australia, and at: southern and western edges of the Gulf of Carpentaria in Queensland and the Northern Territory, Australia has a has light https://www.flickr.com/photos/54876436@N08/9247200620/ grey to mouse brown pelage and with the mid-line from the from Theda Station, Western Australia. snout not going past the crown, being very indistinct and is grey Photos of Pseudochirops (Petropseudes) jamesbondi sp. nov. in in colour, breaking up as it moves up the snout. White under life can be seen online at: forebody. https://www.flickr.com/photos/colonel_007/49215178836/ The white above and below the eye giving an appearance of a from Hells Gate, Queensland, Australia and at: tight circle around the eye is well defined. Inside of ears is https://www.flickr.com/photos/58349528@N02/46592424132/ yellow-orange becoming brownish on the outer edges. from Roper Bar, Northern Territory, Australia. There is no obvious reddish brown or orange overlay on the fur of upper legs save for the same slight and very feint orange Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. from overlay as seen over all parts of the body. Groote Eylandt, Northern Territory, Australia, in life can be seen online at: Pseudochirops (Petropseudes) fiacummingae sp. nov. from the Kimberley district of north-west Western Australia has a light https://denr.nt.gov.au/__data/assets/pdf_file/0003/758307/ grey pelage with the mid line from snout distinct, thin, blackish in groote-archipelago-threatened-species-management-plan-2019- colour and it goes past the crown of the head, without thickening 2028.pdf between the eyes and snout or again before the line between (see page 7) the ears. White under forebody. Document is called The white above and below the eye giving appearance of tight “Groote Archipelago Threatened Species Management Plan circle around eye is not well defined. Inside of ears is orange to 2019-2028” salmon colour with no blackening on the outer edges. Very weak and is a 64 page pdf posted online but has no listed author or reddish brown overlay on fur of upper legs. publisher. Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. from Distribution: Pseudochirops fiacummingae sp. nov. is known Groote Eylandt, Northern Territory, Australia, is similar in most only from the hilly, rocky parts of the Kimberley division of respects to P. jamesbondi sp. nov. except for the obvious and Western Australia, with almost all records in the west and north- strong light orange colour of the rear of the rump and the back west part of the region. legs (versus either not so, or only faint orange tinge in Gulf form, Etymology: Named in honour of one of Australia’s best ever being in line with the rest of the dorsal surface). The mid-line investigative journalists, Fia Cumming of Lyons in the ACT, marking from the snout does not go beyond the crown as in it Australia in recognition for her services to journalism and wildlife terminates well before it and it is grey in colour. conservation in Australia. Further detail can be found in Hoser This marking is noticeably widest at the midpoint between the (1996). eyes, appearing like an elongated diamond shape. Fur on the PSEUDOCHIROPS (PETROPSEUDES) JAMESBONDI SP. lower forelimbs also has a strong orange tinge, but this is only NOV. on the tips, rather than all over as seen on the hind limbs. LSID urn:lsid:zoobank.org:act:B7A2B620-A907-4FB9-982A- All four species of possum in the subgenus Petropseudes ADDAAC5D6DD5 Thomas, 1923 (see above) are readily separated from all other Holotype: A preserved specimen (skeleton and skin) at the possums occurring in the same region in western Queensland, Museum and Art Gallery of the Northern Territory Mammal the Northern Territory and Western Australia by a combination of Collection, specimen number U1155 collected from Echo Gorge, having a much shorter tail, being less than two thirds the length Wollogorang Station, Northern Territory, Australia, Latitude - of the head and body as well as having very dense fur. 17.18 S., Longitude 137.72 E. This government-owned facility Photos of Pseudochirops dahli (Collett, 1895) in life can be seen allows access to its holdings. in Strahan (1988) at page 132 (top) and online at: Paratypes: Two preserved specimens at the Museum and Art https://www.flickr.com/photos/88708273@N03/40093596073/ Gallery of the Northern Territory Mammal Collection, specimen and at: numbers U4186 and U4187 collected from Aquarium Springs Gorge, Wollogorang Station, Northern Territory, Australia, https://www.flickr.com/photos/128365570@N04/42868251355/ Latitude -17.48 S, Longitude 137.63 E. both from Arnhem Land, Northern Territory, Australia. Diagnosis: Until now, all populations of so-called Rock Ringtail Photos of Pseudochirops (Petropseudes) fiacummingae sp. nov. Possum have been assigned to the same putative species, in life can be seen online at: namely Pseudochirops (Petropseudes) dahli (Collett, 1895) with https://www.flickr.com/photos/mattsummerville/48131308123/ a type locality of Mary River. Northern Territory, Australia. from Prince Regent, Western Australia, Australia, and at: That form is now herein restricted to the type locality and https://www.flickr.com/photos/54876436@N08/9247200620/ adjacent hills and outliers, essentially including the Arnhem from Theda Station, Western Australia. Land escarpment and adjacent hills and ranges to the south and west. Photos of Pseudochirops (Petropseudes) jamesbondi sp. nov. in Hoser 2020 - Australasian Journal of Herpetology 42:23-30. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 28 Australasian Journal of Herpetology life can be seen online at: eye is very well defined. Inside of ears is salmon colour with https://www.flickr.com/photos/colonel_007/49215178836/ significant blackening towards the outer edges. from Hells Gate, Queensland, Australia and at ; Pseudochirops (Petropseudes) jamesbondi sp. nov. from the https://www.flickr.com/photos/58349528@N02/46592424132/ southern and western edges of the Gulf of Carpentaria in Queensland and the Northern Territory, Australia has a has light from Roper Bar, Northern Territory, Australia. grey to mouse brown pelage and with the mid-line from the Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. from snout not going past the crown, being very indistinct and is grey Groote Eylandt, Northern Territory, Australia, in life can be seen in colour, breaking up as it moves up the snout. White under online at: forebody. https://denr.nt.gov.au/__data/assets/pdf_file/0003/758307/ The white above and below the eye giving an appearance of a groote-archipelago-threatened-species-management-plan-2019- tight circle around the eye is well defined. Inside of ears is 2028.pdf yellow-orange becoming brownish on the outer edges. (see page 7) There is no obvious reddish brown or orange overlay on the fur Document is called, of upper legs save for the same slight and very feint orange “Groote Archipelago Threatened Species Management Plan overlay as seen over all parts of the body. 2019-2028” Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. from and is a 64 page pdf posted online but has no listed author or Groote Eylandt, Northern Territory, Australia, is similar in most publisher. respects to P. jamesbondi sp. nov. except for the obvious and Distribution: Pseudochirops jamesbondi sp. nov. is known only strong light orange colour of the rear of the rump and the back from the south and west side of the Gulf of Carpentaria, legs (versus either not so, or only faint orange tinge in Gulf form, including nearby hills, where suitable rocky hilly habitat occurs, being in line with the rest of the dorsal surface). The mid-line within far north-west Queensland and adjacent parts of the marking from the snout does not go beyond the crown as in it Northern Territory, Australia. terminates well before it and it is grey in colour. Etymology: Named in honour of James Bond of Park Orchards, This marking is noticeably widest at the midpoint between the (Melbourne), Victoria, Australia in recognition of his role in eyes, appearing like an elongated diamond shape. Fur on the conservation of Australian wildlife, including via his vitally lower forelimbs also has a strong orange tinge, but this is only important work done over many years maintaining the wildlife on the tips, rather than all over as seen on the hind limbs. breeding facility at Snakebusters: Australia’s best reptiles Pseudochirops (Petropseudes) fiacummingae sp. nov. from the shows. Kimberley district of north-west Western Australia has a light PSEUDOCHIROPS (PETROPSEUDES) grey pelage with the mid line from snout distinct, thin, blackish in WADDAMADDAWIDYU SP. NOV. colour and it goes past the crown of the head, without thickening LSID urn:lsid:zoobank.org:act:6644706A-73BE-4861-B175- between the eyes and snout or again before the line between 972349EC27A0 the ears. White under forebody. Holotype: A preserved specimen at the Australian Museum, The white above and below the eye giving appearance of tight Sydney, New South Wales, Australia, specimen number circle around eye is not well defined. Inside of ears is orange to M.21477, collected from the fourth beach near Alyongula, salmon colour with no blackening on the outer edges. Very weak Groote Eylandt, Northern Territory, Australia, Latitude -13.97 S., reddish brown overlay on fur of upper legs. Longitude 136.58 E. This government-owned facility allows All four species of possum in the subgenus Petropseudes access to its holdings. Thomas, 1923 (see above) are readily separated from all other Paratypes: 1/ A preserved female specimen at the Australian possums occurring in the same region in western Queensland, Museum, Sydney, New South Wales, Australia, specimen the Northern Territory and Western Australia by a combination of number M.24782 collected from near Alyangula Creek on the old having a much shorter tail, being less than two thirds the length Rifle Range Road, Groote Eylandt, Northern Territory, Australia, of the head and body as well as having very dense fur. Latitude -13.85 S., Longitude 136.41 E. Photos of Pseudochirops dahli (Collett, 1895) in life can be seen 2/ A preserved specimen (skeleton and skin) at the Museum and in Strahan (1988) at page 132 (top) and online at: Art Gallery of the Northern Territory Mammal Collection, Darwin, https://www.flickr.com/photos/88708273@N03/40093596073/ Northern Territory, Australia, specimen number U3282 collected and at: from Angurugu, Groote Eylandt, Northern Territory, Australia, https://www.flickr.com/photos/128365570@N04/42868251355/ Latitude -13.98 S., Longitude 136.45 E. both from Arnhem Land, Northern Territory, Australia. Diagnosis: Until now, all populations of so-called Rock Ringtail Possum have been assigned to the same putative species, Photos of Pseudochirops (Petropseudes) fiacummingae sp. nov. namely Pseudochirops (Petropseudes) dahli (Collett, 1895) with in life can be seen online at: a type locality of Mary River. Northern Territory, Australia. https://www.flickr.com/photos/mattsummerville/48131308123/ That form is now herein restricted to the type locality and from Prince Regent, Western Australia, Australia, and at: adjacent hills and outliers, essentially including the Arnhem https://www.flickr.com/photos/54876436@N08/9247200620/ Land escarpment and adjacent hills and ranges to the south and from Theda Station, Western Australia. west. Photos of Pseudochirops (Petropseudes) jamesbondi sp. nov. in This species and the three newly described forms herein can be life can be seen online at: readily separated from one another by consistent differences in https://www.flickr.com/photos/colonel_007/49215178836/ colour of pelage on various parts of the head and body. from Hells Gate, Queensland, Australia and at: Nominate P. dahli is readily separated from the other three species by having dark grey pelage and with mid line, from https://www.flickr.com/photos/58349528@N02/46592424132/ snout being distinct, thick, blackish in colour and extends past from Roper Bar, Northern Territory, Australia. the crown of the head. This line becomes wider and darker Pseudochirops (Petropseudes) waddamaddawidyu sp. nov. from between eyes and snout and again before a line between the Groote Eylandt, Northern Territory, Australia, in life can be seen ears. There is a strongly dark reddish brown overlay on fur of the online at: upper legs. Yellowish white under forebody. The white above and https://denr.nt.gov.au/__data/assets/pdf_file/0003/758307/ below the eye, giving the appearance of a tight circle around the groote-archipelago-threatened-species-management-plan-2019-

2028.pdf a further specimen of the type subspecies of P. archeri archeri in (see page 7) life from Ravenshoe, Atherton Tableland, Queensland can be Document is called: seen online at: “Groote Archipelago Threatened Species Management Plan https://www.flickr.com/photos/mattsummerville/16918647658/ 2019-2028” and is a 64 page pdf posted online but has no listed and a further two photos of specimens of the type subspecies of author or publisher. P. archeri archeri in life from the Atherton Tableland, Queensland Distribution: Pseudochirops waddamaddawidyu sp. nov. is can be seen online at: known only from Groote Eylandt in the Northern Territory, https://www.flickr.com/photos/jpmckenna/6547010879/ Australia. Reports of specimens on the immediately proximal and at: Northern Territory mainland and smaller islands to the north are assumed to be of this species. https://www.flickr.com/photos/euprepiosaur/8453294248/ Etymology: On 6 March 2012, I met a Warnindilyakwa Distribution: Pseudochirops (Sloppossum) archeri Aboriginal elder, being the native tribe from Groote Eylandt, who chrismaxwelli subsp. nov. is known only from the Mount Lewis have apparently lived there for some thousands of years. I area of far north Queensland, Australia, being the immediate asked the elderly man for the native name for the local Rock vicinity of the collection localities of the holotype and paratypes. Ringtail Possum to which he replied “waddamaddawidyu”. This is an area of high altitude forests, noting the species generally occurs at elevations in excess of 300 metres above Hence the scientific name. I have since been advised that his sea level and is most commonly seen 500 metres and higher. reply to me actually meant “what is the matter with you?”, because he did not understand the nature of my question, but Etymology: Named in honour of former lawyer and barrister the name has stuck and in the absence of an alternative, this is Christopher Murray Maxwell AC. the scientific name assigned to the taxon. He later succeeded Justice John Winneke as President of the Spelling of the name is intentional and should not be changed. Victorian Court of Appeal on 16 July 2005 and is recognized for PSEUDOCHIROPS (SLOPPOSSUM) ARCHERI his contributions to human rights and justice as well as his fair CHRISMAXWELLI SUBSP. NOV. and equitable application of the law in Australia via his role as a LSID urn:lsid:zoobank.org:act:4ED60895-A1B9-4F02-876E- Court of Appeal judge. F0F278BBB5D7 In the period 1999-2001 he fought valiantly to stop the corrupt Victorian government from outlawing the best-selling books Holotype: A preserved male specimen in the Queensland Victoria Police Corruption (Hoser, 1999a) and Victoria Police Museum, Brisbane, Queensland, Australia, specimen number Corruption - 2 (Hoser 1999b) including an attempt to have JM3617, collected from Mount Lewis, Queensland, Australia, myself (Raymond Hoser) as author, charged with fake criminal Latitude -16.58 S., Longitude 145.26 S. This government-owned charges and potentially jailed indefinitely for exposing endemic facility allows access to its holdings. government corruption (via a so-called “sedition” charge). Paratypes: 1/ A preserved female specimen in the Queensland Incidentally, Maxwell was one of two (out of three) Supreme Museum, Brisbane, Queensland, Australia, specimen number Court Judges who in the Court of Appeal in Victoria upheld a JM9476, collected from Kingfisher Park, Julatten, Queensland, unanimous jury verdict (from 11 December 2018) that Catholic Australia, Latitude -16.62 S., Longitude 145.33 E. Cardinal George Pell had indecently assaulted two Choir boys. 2/ A preserved female specimen in the Queensland Museum, The (alleged) offences occurred in December 1996 and early Brisbane, Queensland, Australia (adult skin and skull), collected 1997 at St Patrick’s Cathedral, months after Pell was from Mossman, Queensland, Australia, Latitude -16.47 S., inaugurated as Archbishop of Melbourne. Longitude 145.38 E. Pell’s Barrister, Robert Richter had sought leniency for his client Diagnosis: Until now Pseudochirops (Sloppossum) archeri Pell on the basis that Pell’s attacks on the young boys was “no chrismaxwelli subsp. nov., from the northern wet tropics region, more than a plain vanilla sexual penetration case” whereby he generally north of Port Douglas and south of Cape Tribulation had forced the boys to suck his penis. including the high elevation and high rainfall hills and plateau Pell sought and got leave to appeal the jury decision to the near the coast has been treated as the same species as Australian High Court (rare in itself) and the case was heard on nominate P. archeri Matschie, 1915, this being the form from the 10 and 11 March 2020, with a decision normally expected to be southern wet tropics region generally north of Mount Spec and handed down several months later. south of Cairns and including the Bellenden Ker Range, However at the height of the Australian Covid-19 Coronavirus Atherton Tableland and Paluma Range. The two subspecies are pandemic, at a time that Australians and the media were separated by the relatively low elevation zone known as the preoccupied with the ongoing deaths and economic disruption Black Mountain Corridor, or alternatively Black Mountain Gap. that accompanied the pandemic, via house lock downs and the P. archeri chrismaxwelli subsp. nov. is readily separated from P. like, the High Court on 7 April rushed out a unanimous decision archeri archeri by having relatively poorly defined markings and stating that the jury had got it wrong and that Pell should have white patches on the face and body (versus bold in P. archeri been acquitted, thereby quashing the conviction. archeri) and the dorsal body fur has a weak greenish tinge, Police did not charge Pell in relation to numerous other alleged versus a strong greenish tinge in P. archeri archeri. sexual assaults of minors. A photo of P. archeri chrismaxwelli subsp. nov. in life from Mount Unfortunately, even when an ethical and highly qualified judge in Lewis, Queensland, can be seen online at: Australia upholds the law, others in the system that is inherently https://www.flickr.com/photos/euprepiosaur/6809527935 corrupt will undermine their excellent work. and at: Meanwhile, in Australia, sex offenders in high places, including https://www.flickr.com/photos/zimny_anders/22113501931/ within the Catholic Church, continue to be corruptly protected by The type subspecies of P. archeri archeri in life from others in the church, police force and judiciary. Yungaburra, Atherton Tableland, Queensland, can be seen CONSERVATION OF THE NEWLY NAMED TAXA online at: In terms of conservation of each population of each species as https://www.flickr.com/photos/bryanjsmitheci/26617682041/ described in this paper, the comments in Hoser (1991 and and at: 2019a, 2019b) apply. https://www.flickr.com/photos/mattsummerville/43431336910/ Wildlife laws as currently enforced in Australia are not in any materially significant way enhancing the long-term survival as well as at: prospects of any of the relevant species and are being vastly Hoser 2020 - Australasian Journal of Herpetology 42:23-30. https://www.flickr.com/photos/mattsummerville/31933820964/ Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 30 Australasian Journal of Herpetology outweighed by other negative impacts of governments, including Diet selection in the green ringtail possum (Pseudochirops their ongoing commitment to growing the human population to a archeri): A specialist folivore in a diverse forest. Austral Ecology level that can only put further pressure on the survival prospects 31(7)799-807. of them. If the Australian government persists with its “Big Laurance, W. 1990. Comparative Responses of five Arboreal Australia Policy”, (see for example Saunders 2019 or Zaczek Marsupials to Tropical Forest Fragmentation. Journal of 2019), then all sorts of unforseen threats to the survival of these Mammalogy, 71:641-653. species will certainly emerge. Lawlor, T. E. 1979. Handbook to the Orders and Families of REFERENCES CITED Living Mammals. Mad River Press, Inc. Eureka, California, USA. Bannister, J. L., Calaby, J. H., Dawson, L. J., Ling, J. K., Maxwell, S., Burbidge, A. A. and Morris, K. 1996. The 1996 Mahoney, J. A., McKay, G. M., Richardson, B. J., Ride, W. D. L. Action Plan for Australian Marsupials and Monotremes. and Walton, D. M. 1988. Zoological Catalogue of Australia, Australasian Marsupial and Monotreme Specialist Group, IUCN Volume 5, Mammals. 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Commission on Zoological Nomenclature) 1999. International Australasian Journal of Herpetology 24:16-19. code of Zoological Nomenclature. The Natural History Museum - Hoser, R. T. 2015. Australian agamids: Eighteen new species Cromwell Road, London SW7 5BD, UK. from the genera Amphibolurus Wagler, 1830, Lophognathus Runcie, M. J. 1999. Biparental care and obligate monogamy in Gray, 1842, Rankinia Wells and Wellington, 1984, Diporiphora the rock-haunting possum, Petropseudes dahli, from tropical Gray, 1842, Tympanocryptis Peters, 1863, as well as three new Australia. Animal Behaviour. 59 (5): 1001-1008. genera and six new subgenera. Australasian Journal of Herpetology 30:37-64. Saunders, S. 2019. Morrison and Shorten’s Big Australia: The overpopulation problem. News article published 17 April 2019 Hoser, R. T. 2016a. 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Catalogue of the Marsupialia and Odatria (Kimberleyvaranus) glebopalma (Mitchell, 1955) Monotremata in the Collection of the British Museum (Natural (Reptilia: Squamata: Varanidae). Australasian Journal of History). British Museum, London, UK:xiii+401 pp+28 pl. Herpetology 37:46-47. Thomas, O. 1922. A selection of lectotypes of the typical Hoser, R. T. 2019a. 11 new species, 4 new subspecies and a Australian marsupials in the British Museum Collection. Ann. subgenus of Australian Dragon Lizard in the genus Mag. Nat. Hist. 9(10):127-128. Tympanocryptis Peters, 1863, with a warning on the Thomas, O. 1923. On some Queensland Phalangeridae. Ann. conservation status and long-term survival prospects of some Mag. Nat. Hist. 9(11):246-250. newly named taxa. Australasian J. of Herpetology 39:23-52. van Ufford, A. Q. and Cloos, M. 2005. Cenozoic tectonics of Hoser, R. T. 2019b. Richard Shine et al. (1987), Hinrich Kaiser et New Guinea. Am. Assoc. Pet. Geol. Bull. 89:119-140. al. (2013), Jane Melville et al. (2018 and 2019): Australian Vaughan, T. A. 1986. Mammalogy, 3rd Edition. Harcourt Brace Agamids and how rule breakers, liars, thieves, taxonomic Jovanovich College Publishers. New York, USA. vandals and law breaking copyright infringers are causing reptile Zaczek, Z. 2019. Former PM Kevin Rudd says Australia should Hoser 2020 - Australasian Journal of Herpetology 42:23-30. species to become extinct. Australasian J. of Herp. 39:53-63. drastically increase its population to 50 MILLION to make sure Iredale, T. and Troughton, E. L. G. 1934. AvailableA check-list of theonline atnation www.herp.net can defend itself in the face of threats from China. Daily mammals recorded from Australia. Australian Museum Memoir Mail (Australia) (27 November), published online at: https:// 6:1-122. [4 May 1934]. Copyright- Kotabi Publishingwww.dailymail.co.uk/news/article-7729689/Kevin-Rudd-says- - All rights reserved Jones, K. W., Maclagen, S. J. and Krockenberger, A. K. 2006. Australia-increase-population-50-MILLION-defend-China.html CONFLICTS OF INTEREST - NONE Australasian Journal of Herpetology 31 Australasian Journal of Herpetology 42:31-37. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

Small, easily overlooked and in decline. Potoroos in Eastern Australia. A formal division of the genus Potorous Desmarest, 1804 (Marsupialia Potoroidae) and the description of a new species from south-east Queensland.

LSID URN:LSID:ZOOBANK.ORG:PUB:339C5641-9CA7-41A3-9436-1DADAF52B67B

RAYMOND T. HOSER

LSID URN:LSID:ZOOBANK.ORG:AUTHOR:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 26 February 2020, Accepted 23 March 2020, Published 25 April 2020.

ABSTRACT Potoroos within the family Potoroidae are small marsupials which were abundant at the time Europeans first came to Australia (White and Stone 1790). They have severely declined in number since. Three main species groups, all currently placed in the genus Potorous Desmarest, 1804 (type species Didelphis tridactyla Kerr, 1792) have been formally described and named, containing one putative species each based on most recently published classifications. One of these, the Long-nosed Potoroo Potorous tridactylus (Kerr, 1792) was recently subdivided into two species, namely P. gilberti (Gould, 1841) from Western Australia and P. tridactylus from eastern Australia, treated as consisting three subspecies, being the nominate form from the Central Coast of New South Wales, P. tridactylus apicalis (Gould, 1851) from Tasmania and P. tridactylus trisulcatus (McCoy, 1865) from Victoria (Frankham et al. 2012). Molecular studies have shown east Australian P. tridactylus to consist of four main divergent clades and so the unnamed one is formally named in this paper as P. waddahyamin sp. nov. based on well-known morphological divergences. Each of the four clades are also formally elevated to full species based on known dates of divergence being 1.32 and 2.45 MYA from nearest common ancestor. Of the three main species groups within the putative genus Potorous two have generic names available being Potorous and Potoroops Matschie, 1916 for the type species Hypsiprymnus platyops Gould, 1844. The third species group does not. As the molecular studies of Westerman et al. (2004) and Frankham et al. (2012) showed genus-level divergences between the groups, the unnamed one is formally named for the first time. The species Potorous longipes Seebeck and Johnston, 1980 is formally placed in the new genus Rossignolius gen. nov.. Keywords: Taxonomy; nomenclature; classification; Potoroo; Marsupials; Potoroidae; Potorous; Potoroops; Hypsiprymnus; platyops; tridactylus; trisulcatus; gilberti; apicalis; longipes; New genus; Rossignolius; new species; waddahyamin.

INTRODUCTION (Macropodidae). Potoroos exhibit many morphological Potoroos within the family Potoroidae are (now) rarely seen specializations such as an elongated pointed rostral region, rabbit-sized marsupials which were abundant at the time erect ears, large eyes, large claws for digging, and long robust Europeans first came to Australia (White and Stone 1790). hind legs. At first glance, the animal with its pointed nose and grey-brown The body length varies between species, but is usually between fur looks very much like a better-known bandicoot (family 34 and 38 cm and the semi-prehensile tail length is 15 to 24 cm. Peramelidae), until it hops away with its front feet tucked into its As they are nocturnal, usually live in forests and are rarely seen chest, revealing its closer relationship with the kangaroo family in the wild, except as road kill, better indicators of its presence Hoser 2020 - Australasian Journal of Herpetology 42:31-37. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 32 Australasian Journal of Herpetology are the runways it makes through the undergrowth and the in line with what has just been said and that P. longipes was in hollow diggings it leaves behind when feeding on underground need of a genus-level reassignment. roots and fungi. Before doing any of the above, it was however important to Due to the introduction by Europeans of competing animals like review the relevant literature as well as inspect specimens of all rabbits and hares and predators like foxes, cats (see Spencer relevant putative taxa. 1991) and dogs, numbers of Potoroos severely declined almost MATERIALS, METHODS AND RESULTS immediately after European settlement in the 1700’s. Before a decision is made to name any new taxon, reasonable These animals only remain in large numbers on islands that are steps must be taken to ensure that it is justified on all relevant effectively predator free and on the east coast in relatively grounds, including that it is morphologically, genetically and undisturbed forest areas, which in effect also form islands of reproductively isolated from their nearest relative and to a safety from invasive species that do better in disturbed and sufficient degree to be of taxonomic significance. agricultural habitats. A further relevant question to ask is should the reproductively The type species for the genus is the Long-nosed Potoroo isolated and morphologically divergent entities be labelled as Potorous tridactylus (Kerr, 1792), originally named Didelphis subspecies, full species, or potentially higher level again. tridactyla Kerr, 1792. The genus Potorous was erected by Key literature relevant to the taxonomic and nomenclatural Matschie in 1916. This was a widespread putative taxon, being conclusions within this paper include Abbott (2008), Amos found in south-west Australia and the south-east as far north as (1982), Bannister et al. (1988), Bee and Close (1993), Bennett wetter parts of south-east Queensland. (1993), Bensley (1903), Bougher and Friend (2009), Browning et The south-west Australian form was named Hypsiprymnus al. (2001), Bryant and Krosch (2016), Burbidge and Manly gilberti by Gould in 1841, based on a type from King George (2002), Butler and Merrilees (1970), Byrne et al. (2008, 2011), Sound, but later synonymised with P. tridactylus by Iredale and Claridge et al. (2007, 2010), Courtney (1963), Desmarest Troughton (1934) and Bannister et al. (1988). (1804), Dexter and Murray (2009), Eldridge (1997), Eldridge and Likewise for the putative taxa Hypsiprymnus apicalis Gould, Close (1992), Eldridge and Potter (2020), Eldridge et al. (2001), 1851 from New Norfolk, Tasmania and H. trisulcatus McCoy, Finlayson (1938), Ford (2014), Frankham et al. (2011, 2012), 1865 based on a type from near Gisborne, Victoria. Friend (2003), Groves et al. (2005), Guiler (1958, 1960, Guiler Frankham et al. (2012) published a phylogeny for the putative and Kitchener (1967), Gould (1841, 1844, 1851), Gray (1837), species P. tridactylus including specimens from across the Heinsohn (1936), Hoke (1990), Hope (1969), Hoser (1991), known range and found significant divergences between the Iredale and Troughton (1934), Jackson (2008), Johnson (2003), populations. Johnston (1973), Johnston and Sharman (1976, 1977), Johnston et al. (1984), Kerr (1792), Kitchener (1973), Lawlor They found putative P, gilberti diverged from the (south) eastern (1979), Linné et al. (1792), Long (2001), Mahoney (1964), populations 5.77 MYA and 7.2 MYA from those from the New Mason (1997), Matschie (1916), Maxwell et al. (1996), McCoy South Wales (NSW) Central Coast and further north (a total of (1865), McDowell and Medlin (2010), Menkhorst (2001), two main identified groups). In turn they found Tasmanian Meredith et al. (2008), Mitchell (1908), Nguyen et al. (2005), specimens diverged from those of Victoria and southern NSW Norton et al. (2010a, 2010b), Potter et al. (2012a, 2012b, 2012c, by 2,45 MYA and the two northern populations to have diverged from one another 1.32 MYA. 2014), Ride (1970), Ride et al. (1999), Rounsevell et al. (1991), Seebeck (1991), Seebeck and Johnston (1980), Shaw (1800), They made the taxonomic judgements of recognizing P, gilberti Short (1998), Shortridge (1910), Sinclair and Westerman (1997), as a separate species and in turn recognizing the single species Sinclair et al. (1996, 2000, 2002), Spencer (1991), Stead- P. tridactylus, with both P. apicalis and P. trisulcatus as Richardson et al. (2010), Strahan (1988), Tate (1948), Thomas subspecies of that. (1888, 1909), Vaughan et al. (2007), Vaughan (1986), Vaughan- The taxonomy did not make sense in that both putative species Higgins et al. (2011), Vernes and Jarman (2014), Westerman et P. apicalis and P. trisulcatus (diverged from one another 2.45 al. (2004, 2010), White and Stone (1790) as well as sources MYA) were in fact more closely related to putative P, gilberti, with cited therein (duplicitous references not necessarily included). a 5.77 MYA divergence as opposed to a 7.2 MYA divergence from putative P. tridactylus. Live and dead specimens as well as available bone specimens, In order to resolve this clear error of assignment, one must were examined as was other relevant material, including past either transfer both putative subspecies to the synonymy of P, climate data for the applicable regions, sea level depths and gilberti and continue to regard each as subspecies, or in the other relevant information. alternative, and based on quoted divergences, simply elevate all In summary, as inferred already, the results of Frankham et al. to full species, which in the absence of evidence to the contrary (2012) as interpreted and corrected in the introduction stood up would be the most sensible outcome. to scrutiny. Therefore P. tridactylus as currently recognized is herein split In terms of the remainder of putative P. tridactylus, Frankham et into four species, as well as the sometimes separately al. (2012) found a divergence between two main groups of 1.32 recognized species P. gilberti, making a total of five. MYA, which is again a species-level divergence. The unnamed taxon from south-east Queensland, with a With the northern group unnamed, it again seemed logical to divergence of 1,32 MYA from nearest congener is formally name that as a new species to enable proper conservation named P. waddahyamin sp. nov.. measures to take place and allow zoologists to do proper studies involving known taxa as opposed to directly transposing I note that a 1.32 MYA is definitely worthy of taxonomic potentially misleading results from other taxa to it. recognition and that as recently as year 2020, Eldridge and Potter (2020), gave taxonomic recognition to a central Australian In terms of the morphologically similar species formally named population of Rock Wallaby with a divergence less than 1 MYA P. longipes Seebeck and Johnston, 1980 from south-east from the nominate form. Australia, Frankham et al. (2012) found a divergence of 9.4 MYA which is a genus-level divergence. Potorous Desmarest, 1804 with a type species of Didelphis tridactyla Kerr, 1792 has until now been treated by most Westerman et al. (2004) found the putative species P. platyops publishing authors as having three putative species. (Gould, 1844) to have diverged 14.5 MYA from nearest common These are P. tridactylus, herein divided into five as outlined ancestor of the other extant putative Potorous species. above as well as two others. Hence on the basis of the preceding, it seemed self-evident that the taxonomy of the P. tridactylus group needed to be reviewed The other species are Hypsiprymnus platyops Gould, 1844, better-known as the Broad-faced Potoroo, believed to be extinct

(Hoser, 1991), and the Long-footed potoroo Potorous longipes 2150 (from the present 25 million as of 2019), all sorts of Seebeck and Johnston, 1980. unforseen threats to the survival of these species may emerge. Westerman et al. (2004) found the putative species P. platyops ROSSIGNOLIUS GEN. NOV. (Gould, 1844) to have diverged 14.5 MYA from the nearest LSID urn:lsid:zoobank.org:act:2D06792C-B5AE-43D3-9A9D- common ancestor of the other extant putative Potorous species. FF1F16C77661 Hence, after taking the divergent morphology of the taxon into Type Species: Potorous longipes Seebeck and Johnston, 1980. account, there is no doubt it is highly divergent of the other two Diagnosis: Rossignolius gen. nov. has until now been treated species. The genus name Potoroops Matschie, 1916 is available as within the genus Potorous Desmarest, 1804 but is readily for this taxon and so is adopted herein in line with Tate (1948) separated from all members of that genus and the based on the significantly divergent morphology as detailed by morphologically similar Potoroops Matschie, 1916 by the Tate (1948). following unique suite of characters: larger adult size, with an Frankham et al. (2012) in their published phylogeny of the other average 400 mm head and body length, average 320 mm tail living species found a 9.4 MYA divergence between P. length, 2.1 kg average weight of males and 1.7 kg average tridactylus (all five forms adopted as species herein) and weight of females (versus .9 to 1.1 kg in Potorous and .5 kg in Potorous longipes Seebeck and Johnston, 1980, which is clearly Potoroops, both sexes of each); grey brown above and grey a divergence worthy of genus-level separation. below; hind foot is longer than the head (versus not so in I note other mammals have been divided into separate genera Potorous); the skull is lightly built, long and narrow, (versus on the basis of far lower divergences than this including for broad in Potoroops); there is a distinctive pad on the sole of the example Petrogale Gray, 1837 and Peradorcas Thomas, 1909 first (big) toe; chromosome count of 34 (versus 12 in males and with a divergence estimated at under 9 MYA (Potter et al. 13 in females in Potorous). 2012b). The genus Potorous Desmarest, 1804, sensu lato including the The species Potorous longipes Seebeck and Johnston, 1980 is species within Rossignolius gen. nov. and Potoroops Matschie, therefore formally placed in the new genus Rossignolius gen. 1916 are readily separated from other marsupials by the nov.. following suite of characters: Rhinarium naked, variable in its INFORMATION RELEVANT TO THE FORMAL DESCRIPTIONS extent on the upperside of the muzzle. Ears very short, rounded. THAT FOLLOW Fore claws long, rather slender, the median much longer than the outer ones. Hind feet very short, the whole hind limb not There is no conflict of interest in terms of this paper or the disproportionally longer than the fore; soles naked and coarsely conclusions arrived at herein. granulated. Tail shorter than in the morphologically similar Several people including anonymous peer reviewers who Bettongia, tapering, covered with short closely adpressed hairs revised the manuscript prior to publication are also thanked as and without any trace of a crest. are relevant staff at museums who made specimens and records available in line with international obligations. Within the genus Potorous Desmarest, 1804, sensu stricto the skull is lightly built, long and narrow, the degree of this varying In terms of the following formal descriptions, spellings should between species being relatively shortest in Potorous not be altered in any way for any purpose unless expressly and waddahyamin sp. nov. formally described in this paper. In the exclusively called for by the rules governing Zoological genus Potoroops Matschie, 1916 the skull is distinctively and Nomenclature as administered by the International Commission obviously short as well as being very wide. The interorbital of Zoological Nomenclature. space is broad, flat or slightly convex, its edges square, not Material downloaded from the internet and cited anywhere in this ridged, but in old age forming short postorbital processes. paper as being sourced online was downloaded and checked Anterior palatine foramina very short. Palate with a single pair of most recently as of 22 January 2020 (including if also viewed rather small vacuities opposite the posterior molars, and entirely prior), unless otherwise stated and was accurate in terms of the confined to the palatal bone. Bullae very slightly swollen to content (as described) cited herein as of that date. distinctly swollen, nearly flat. Lower jaw long and slender, not Unless otherwise stated explicitly, colour and other descriptions strongly convex below. apply to living adult specimens of generally good health and not There are typically about 34 teeth in the mouth. Upper incisors under any form of stress by means such as excessive cool, small and light, with incisor one long and pointed. Canines are heat, dehydration or abnormal skin or fur reaction to chemical or small and thin, compressed laterally; premolar 4 is long and other input. trenchant, though considerably shorter than in Bettongia, without Unless stated otherwise, colour descriptions of species refer to trace of internal ledge or postero-internal talon, its outer and fur colour and not skin. inner surfaces with from two to four shallow vertical grooves. While numerous texts and references were consulted prior to Decidnous premolar number three is similar but smaller. Molars publication of this paper, the criteria used to separate the quadrangular and quadritubercular, evenly but slightly relevant species or subspecies has already been spelt out and/ decreasing in size backwards, except that molar one is generally or is done so within each formal description and does not rely on rather smaller than molar two. Lower incisors very long and material within publications not explicitly cited herein. slender. In terms of conservation of each population of each species or These animals stand on their hind legs like Bettongs, but unlike subspecies as described below, the comments in Hoser (1991, the Bettongs they run in a different way. 1993, 1996, 2019a and 2019b) apply. These species use the fore as well as the hind legs in a sort of Wildlife laws as currently enforced in Australia are not in any gallop; they also never attempt to kick with their hind legs. materially significant way enhancing the long-term survival A colour photo of a live adult specimen of the only known prospects of any of the relevant species and in any event are species in the genus, namely Rossignolius longipes (Seebeck being vastly outweighed by other negative impacts of and Johnston, 1980) can be found on page 206 of Hoser (1991). governments, including their ongoing commitment to growing Distribution: Known only from a limited area in far north-east the human population to a level that can only put further Victoria and immediately adjacent south-east New South Wales, pressure on the survival prospects of the relevant taxa. in an area closely bound by the coastline (generally within 100 If the Australian government persists with its ecologically km) bounded by Bondi State Forest in New South Wales as the reckless “Big Australia Policy”, (see for example Saunders 2019 north-eastern limit, Orbost, Victoria in the South-west, and or Zaczek 2019), that being a long-term aim to increase the another known population at Howitt Plains, Victoria in the west. human population in Australia to over 100 million people by year Etymology: Rossignolius gen. nov. is named in honour of Hoser 2020 - Australasian Journal of Herpetology 42:31-37. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 34 Australasian Journal of Herpetology

Federico (AKA Fred) Rossignoli of Hurstbridge (Melbourne) in the morphologically similar Bettongia, tapering, covered with Victoria in recognition of his services to herpetology for more short closely adpressed hairs and without any trace of a crest. than two decades, including providing logistical assistances to The skull is lightly built, long and narrow, except in the genus Snakebusters: Australia’s best reptiles shows at times of need Potoroops Matschie, 1916 where it is distinctively short. The and other vitally important services to others involved in Interorbital space is broad, flat or slightly convex, its edges scientific research and wildlife conservation. square, not ridged, but in old age forming short postorbital Through his wildlife display business “Australian Nature Live” processes. Anterior palatine foramina very short. Palate with a Federico educated countless Australians at schools, universities single pair of rather small vacuities opposite the posterior and major public events about reptiles and other wildlife as well molars, and entirely confined to the palatal bone. Bullae very as other public interest matters with some of the most slightly swollen, nearly flat or obviously swollen. Lower jaw long informative and educational shows ever performed. While he and slender, not strongly convex below. has retired from doing these displays as of 2020, methods he There are typically about 34 teeth in the mouth. pioneered are now used by other wildlife displayers worldwide, Upper incisors small and light, with incisor one long and pointed. including by Snakebusters: Australia’s best reptiles shows. Canines are small and thin, compressed laterally; premolar 4 is POTOROUS WADDAHYAMIN SP. NOV. long and trenchant, though considerably shorter than in LSID urn:lsid:zoobank.org:act:0148A5E4-79D6-4270-B331- Bettongia; without trace of internal ledge or postero-internal D1F9EE317B2D talon, its outer and inner surfaces with from two to four shallow Holotype: A preserved male specimen at the Australian vertical grooves. Decidnous premolar number three is similar but Museum in Sydney, New South Wales, Australia, specimen smaller. Molars quadrangular and quadritubercular, evenly but number M.44614.001 collected from near Warwick, Queensland, slightly decreasing in size backwards, except that molar one is Australia, Latitude -28.21 S. Longitude 152.04 E. This generally rather smaller than molar two. Lower incisors very long government-owned facility allows access to its holdings. and slender. Paratypes: Two preserved male specimens at the Australian These animals stand on their hind legs like Bettongs, but unlike Museum in Sydney, New South Wales, Australia, specimen the Bettongs they run in a different way. numbers M.44612.006 and M.44615.002 collected from near These species use the fore as well as the hind legs in a sort of Warwick, Queensland, Australia, Latitude -28.21 S. Longitude gallop; they also never attempt to kick with their hind legs. 152.04 E. A colour photo of a live adult specimen of the only known Diagnosis: Potorous waddahyamin sp. nov. has until now been species in the genus Rossignolius. namely Rossignolius treated as a northern population of the better known P. longipes (Seebeck and Johnston, 1980) can be found on page tridactylus (Kerr, 1792), since subdivided by Frankham et al. 206 of Hoser (1991). (2012) without morphological diagnosis and merely adopting Hoser (1991) at page 205 has a colour reproduction of an image previously proposed and available names. of two Potoroops platyops (Gould, 1841) taken from a lithograph Potorous waddahyamin sp. nov. is readily separated from all of in Gould’s the Mammals of Australia. P. tridactylus (Kerr, 1792), P. apicalis (Gould, 1851), P. The genus Rossignolius gen. nov. has until now been treated as trisulcatus (McCoy, 1865) and P. gilberti (Gould, 1841), by the within the genus Potorous Desmarest, 1804. This genus is following suite of characters: colouration is usually a light brown readily separated from all members of that genus and the pelage with numerous lighter tipped or whitish hairs scattered morphologically similar Potoroops Matschie, 1916 by the throughout giving a distinctly peppered appearance, versus dark following unique suite of characters: Larger adult size, with an brown in the type form of P. tridactylus and dark, slightly reddish average 400 mm head and body length, average 320 mm tail brown in P. trisulcatus and either rufous brown or grey brown in length, 2.1 kg average weight of males and 1.7 kg average P. apicalis; a lack of a white tip on the end of the tail, versus weight of females (versus .9 to 1.1 kg in Potorous and .5 kg in sometimes present in P. tridactylus and P. trisulcatus and Potoroops, both sexes of each); grey brown above and grey usually present in P. apicalis; nasals are short and strongly below; hind foot is longer than the head (versus not so in expanded posteriorly (versus not so in all of P. tridactylus, P. Potorous); the skull is lightly built, long and narrow, (versus apicalis, P. trisulcatus and P. gilberti) and the general fronto- broad in Potoroops); there is a distinctive pad on the sole of the nasal outline is not distinctly and evenly convex as seen in P. first (big) toe; chromosome count of 34 (versus 12 in males and gilberti; skull is relatively shorter in P. waddahyamin sp. nov. 13 in females in Potorous). than all of P. tridactylus; P. apicalis; P. trisulcatus and P. gilberti The genus Potoroops Matschie, 1916 as defined in this paper, is and unlike those species is noticeably expanded posteriorly; readily separated from both Potorous and Rossignolius gen. snout is relatively short in P. waddahyamin sp. nov. as compared nov. by the following unique suite of characters: Naked part of to all of P. tridactylus; P. apicalis; P. trisulcatus and P. gilberti, rhinarium not running back along the top of the muzzle. Head being longest in P. apicalis; P. waddahyamin sp. nov. has a deep short and broad. Hind foot less than 65 mm long. Muzzle very muzzle, versus not so in all of P. tridactylus; P. apicalis; P. short. Orbit to anterior nares 26 mm or less, not exceeding three trisulcatus and P. gilberti; bullae very slightly swollen, versus fourths of the zygomatic breadth. Premolar four is very small, distinctively swollen in P. gilberti and the average weight of about 4-5 mm long. adults of both sexes is just over 1 kg in P. waddahyamin sp. nov. versus under 1 kg in all other species. A photograph of P. waddahyamin sp. nov. in life can be seen online at: The five species in the genus Potorous Desmarest, 1804 as https://imagelibrary.qm.qld.gov.au/fotoweb/archives/5020- defined herein are readily separated from species in the genera Learning-Platform/Icemedia/Mammals/pq019.jpg.info Potoroops Matschie, 1916 and Rossignolius gen. nov. where the relatively short and deep snout of this taxon (as The genus Potorous Desmarest, 1804, sensu lato, including the compared to P. tridactylus; P. apicalis; P. trisulcatus and P. species within Rossignolius gen. nov. and Potoroops Matschie, gilberti) is clearly visible. 1916 are readily separated from other marsupials by the Distribution: P. waddahyamin sp. nov. is known from the coast, following suite of characters: ranges and nearby elevated slopes of south-east Queensland, Rhinarium naked, variable in its extent on the upperside of the south of the dry zone north the Sunshine Coast to near the New muzzle. Ears very short, rounded. Fore claws long, rather South Wales and Queensland border, potentially just into slender, the median much longer than the outer ones. Hind feet northern New South Wales north of the Clarence River. very short, the whole hind limb not disproportionally longer than Etymology: On 30 July 2019, while doing fieldwork on the the fore; soles naked and coarsely granulated. Tail shorter than outskirts of Warwick in south-east Queensland, I asked a local Hoser 2020 - Australasian Journal of Herpetology 42:31-37. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 35

Keinjan Aboriginal elder who spoke the Wakka Wakka language Collingwood, Victoria, Australia. what they called the local Potoroo species, to which he replied Claridge, A. W., Paull, D. J. and Barry, S. C. 2010. Detection of “Waddahyamin”. Hence the scientific name. I was later advised medium-sized ground-dwelling mammals using infrared digital the man may have been confused and had simply asked “What cameras: an alternative way forward? Australian Mammalogy do you mean?” and that there may be no local Aboriginal name 32:165-171. for his animal, so I decided to retain use of “waddahyamin”. The Courtney, J. 1963. The remarkable faunal unit (with description spelling of the scientific name is intentional and should not be of a new form of Potorous rat-kangaroo). Australian Aviculture changed unless mandated by the rules of zoological 17:18-20. nomenclature. Desmarest, A. G. 1804. 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Phylogenetic Zaczek, Z. 2019. Former PM Kevin Rudd says Australia should relationships within the Genus Potorous (Marsupialia: drastically increase its population to 50 MILLION to make sure Potoroidae) based on allozyme electrophoresis and sequence nation can defend itself in the face of threats from China. Daily analysis of the Cytochrome b Gene. Journal of Mammalian Mail (Australia) (27 November), published online at: https:// Evolution. 4(3):147-161. www.dailymail.co.uk/news/article-7729689/Kevin-Rudd-says- Australia-increase-population-50-MILLION-defend-China.html Sinclair, E. A., Danks, A., Wayne, A. F. 1996. Rediscovery of Gilbert’s Potoroo, Potorous tridactylus, in Western Australia. CONFLICTS OF INTEREST Australian Mammalogy. 19(1):69-72. None. Sinclair, E. A., Murch, A. R., Di Renzo, M. and Palermo, M. 2000. Chromosome morphology in Gilbert’s potoroo, Potorous Australasian Journal of gilbertii (Marsupialia: Potoroidae). Aust. J. Zool. 48:281-287. Sinclair, E. 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Monitoring reproduction in the critically endangered marsupial, Gilbert’s potoroo (Potorous gilbertii): preliminary http://www.herp.net analysis of faecal oestradiol-17beta, cortisol and progestagens. Copyright. All rights reserved. General and Comparative Endocrinology.165(1):155-62. Australasian Journal of Herpetology is also a Strahan, R. (ed.) 1988. The Australian Museum Complete book registered trademark ® in all relevant areas and of Australian Mammals / The National Photographic Index of jurisdictions (Australian trademark number: Australian Wildlife. Angus and Robertson, North Ryde, NSW, 1686575). Australia:530 pp. All Intellectual Property (IP) rights are reserved, Tate, G. H. H. 1948. Studies on the anatomy and phylogeny of including in relation to all IP generated by the the Macropodidae (Marsupialia). Bulletin of the American journal in terms of both authors, publisher and the Museum of Natural History 91(2). like.

Hoser 2020 - Australasian Journal of Herpetology 42:31-37. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 38 Australasian Journal of Herpetology Australasian Journal of Herpetology 42:38-41. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

A new subspecies of Yellow-bellied Glider (Marsupialia: Petauridae) from far north Queensland, Australia.

LSID URN:LSID:ZOOBANK.ORG:PUB:836EDBB0-1DD2-42BC-ACB1-FB2D6A6BA199

RAYMOND T. HOSER

LSID urn:lsid:zoobank.org:author:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 6 April 2020, Accepted 16 April 2020, Published 25 April 2020.

ABSTRACT The taxonomy of the east Australian Yellow-bellied Glider Petaurus australis Shaw, 1791 has been confused, conflicting and subject to change over the past 200 years. Several type specimens have been formally described. The nominate form is from New South Wales as are at least three other types, all formally synonymised by Bannister et al. (1988). The putative subspecies Petaurus australis reginae Thomas, 1923 was found by Brown et al. (2006) to be genetically similar to the nominate form and so has been treated as synonymous ever since. However Brown et al. (2006) also showed that the putative P. australis from the wet tropics of far north Queensland, separated from southern populations by the Biogeographic barrier of the Burdekin Gap, are sufficiently divergent both genetically and morphologically to be treated as a subspecies. This view has been agreed by others (Anonymous 2008, Department of the Environment and Resource Management 2011, 2017) and yet the taxon has until now been unnamed. This paper corrects this situation and formally names the subspecies P. australis adelynhoserae subsp. nov.. With an extant population estimated at just 6,000 individuals (Anonymous 2008) and significant ongoing threats causing a long-term decline in the taxon, it is critically important that a sound conservation and captive-breeding program be commenced. Keywords: Mammals; Marsupial; Petauridae; taxonomy; nomenclature; possum; yellow-bellied glider; wet tropics; Queensland; Australia; Petaurus; australis; new subspecies; adelynhoserae.

INTRODUCTION The Yellow-bellied Glider Petaurus australis Shaw, 1791, an Brown et al. (2006) were aware of this fact and subjected this inhabitant of wet sclerophyll forests of eastern Australia, is a form from south-east Queensland and the type form of P. gliding possum with grey-brown or black fur on the upper body australis from New South Wales to a rigorous molecular and a distinctive off-white to yellow or orange belly. It has a fluffy analysis. tail growing to 48 cm long and large, bare ears. Its head and The putative subspecies Petaurus australis reginae Thomas, body grows to 30 cm long and can weigh up to 700 g (Cronin, 1923 was found by Brown et al. (2006) to be genetically similar 1991; Strahan, 2008). to the nominate form and so has been treated as synonymous The taxonomy of Petaurus australis Shaw, 1791 has been ever since. confused, conflicting and subject to change over the past 200 However Brown et al. (2006) also showed that the putative P. years. australis from the wet tropics of far north Queensland, separated Several type specimens have been formally described. The from southern populations by the Burdekin Gap, are sufficiently nominate form is from New South Wales as are at least three divergent both genetically and morphologically to be treated as a other types, all formally synonymised by Bannister et al. (1988). subspecies. Some authors including for example Strahan (1988) mistakenly This view has been agreed by others (Anonymous 2008, assigned the north Queensland population of Petaurus australis Department of the Environment and Resource Management to the subspecies Petaurus australis reginae Thomas, 1923. 2011, 2017) and yet the taxon has until now been unnamed. However that putative subspecies was in fact described from a This paper corrects this situation and formally names the type specimen from Gin Gin in south-east Queensland. subspecies P. australis adelynhoserae subsp. nov.. Hoser 2020 - Australasian Journal of Herpetology 42:38-41. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 39

With an extant population estimated at just 6,000 individuals While numerous texts and references were consulted prior to (Anonymous 2008) and significant ongoing threats causing a publication of this paper, the criteria used to separate the long-term decline in the taxon, including an ever expanding relevant species or subspecies has already been spelt out and/ Australian human population (Hoser, 1991), it is critically or is done so within each formal description and does not rely on important that a sound conservation and captive-breeding material within publications not explicitly cited herein. program be commenced. In terms of conservation of the population of the relevant Hoser (1991) at page 222 also detailed further steps required to subspecies as described below, the comments in Hoser (1991 save the putative species P. australis across its range. and 2019a, 2019b) apply. MATERIALS, METHODS AND RESULTS Wildlife laws as currently enforced in Australia are not in any Before a decision is made to name any new candidate taxon, materially significant way enhancing the long-term survival reasonable steps must be taken to ensure that it is justified on prospects of any of the relevant subspecies and are being vastly all relevant grounds, including that it is morphologically, outweighed by other negative impacts of governments. genetically and reproductively isolated from their nearest relative This includes the Australian National and Queensland State and to a sufficient degree to be of taxonomic significance. governments ongoing commitment to growing the human A further relevant question to ask is, should the reproductively population to a level that can only put further unsustainable isolated and morphologically divergent entities be labelled as pressure on the survival prospects of the relevant subspecies. subspecies, full species, or potentially higher level again. If the Australian government persists with its “Big Australia Key literature relevant to the taxonomic and nomenclatural Policy”, (see for example Saunders 2019 or Zaczek 2019), that conclusions within this paper include being a long-term aim to increase the human population in Anonymous (2008), Bannister et al. (1988), Brown et al. (2004, Australia to over 100 million people by year 2150 (from the 2006), Bryant and Krosch (2016), Collins (1973), Craig (1985), present 25 million as of 2019), all sorts of unforseen threats to Goldingay (1990), Goldingay and Kavanagh (1992), Groves et the survival of this subspecies will almost certainly emerge. al. (2005), Hedges (2006), Hoser (1991), Iredale and Troughton PETAURUS AUSTRALIS ADELYNHOSERAE SUBSP. NOV. (1934), Lawlor (1979), Maxwell et al. (1996), Menkhorst (2001), LSIDurn:lsid:zoobank.org:act:2BA93A74-FB83-4AA6-A464- Ride (1970), Russell (1980), Strahan (1998), Tate (1952), 90A64F7980F8 Thomas (1888, 1922, 1923), Vaughan (1986), Winter (1997, Holotype: A preserved male specimen at Queensland Museum, 2004), Winter et al. (1979) and sources cited therein (duplicitous Brisbane, Queensland, Australia, specimen number JM8746 references not necessarily included). collected from Nichaga Creek, Queensland, Australia, Latitude - Live and dead specimens as well as available bone specimens, 17.82 S., Longitude 145.56 E. This government-owned facility were examined as was other relevant material, including past allows access to its holdings. climate data for the relevant regions, sea level depths, and other Paratypes: Two preserved specimens at the Queensland potentially useful information. Museum, Brisbane, Queensland, Australia, specimen number In summary, as inferred already, the genetic, geological, JM8747 (a male) collected from Nichaga Creek, Queensland, historical and morphological evidence clearly showed that the Australia, Latitude -17.82 S., Longitude 145.56 E. and specimen north Queensland population of P. australis warrants recognition number JM8503 (a female) collected from the Nichaga Creek as a subspecies in accordance with the International Code of Catchment, Queensland, Australia, Latitude -17.83 S., Zoological Nomenclature (Ride et al. 1999). Longitude 145.55 E. For the record, Brown et al. (2006) wrote: “Taking into account Diagnosis: Petaurus australis adelynhoserae subsp. nov. has other behavioural and ecological data, and the disjunct until now been treated (taxonomically at least) as a northern distribution of NQ populations from southern populations, we population of either P. australis australis Shaw, 1791 or propose that the NQ population represents a distinct alternatively the subspecies P. australis reginae Thomas, 1923, Evolutionarily Significant Unit, a lineage showing highly (type locality of Gin Gin, south-east Queensland), now also restricted gene flow from the rest of the species.” treated as a junior synonym of P. australis australis. INFORMATION RELEVANT TO THE FORMAL DESCRIPTION Petaurus australis adelynhoserae subsp. nov. from the wet THAT FOLLOWS tropics region of Northern Queensland, north of the Burdekin There is no conflict of interest in terms of this paper or the Gap (Townsville area) is the taxon erroneously previously conclusions arrived at herein. labelled P. australis reginae by Strachan (1988) and as an undescribed form by others including (Anonymous 2008, Several people including anonymous peer reviewers who Department of the Environment and Resource Management revised the manuscript prior to publication are also thanked as 2011, 2017). are relevant staff at museums who made specimens and records available in line with international obligations. Contrary to the statement on page 312 of Brown et al. (2016), In terms of the following formal description, spelling should not the subspecies Petaurus australis adelynhoserae subsp. nov. is be altered in any way for any purpose unless expressly and readily separated from nominate P. australis australis by being exclusively called for by the rules governing Zoological lighter in pelage, not darker as stated by Brown et al. (2016). Nomenclature as administered by the International Commission Petaurus australis adelynhoserae subsp. nov. is separated from of Zoological Nomenclature. P. australis australis by having a medium to dark brown pelage Material downloaded from the internet and cited anywhere in this on the upper surfaces, versus blackish in P. australis australis. paper as being sourced online was downloaded and checked Petaurus australis adelynhoserae subsp. nov. is further most recently as of 10 February 2020 (including if also viewed separated from P. australis australis by having ill-defined darker prior), unless otherwise stated and was accurate in terms of the fur on the cheeks, versus well defined in P. australis australis. content (as described) cited herein as of that date. The upper fur of the anterior of the tail in Petaurus australis adelynhoserae subsp. nov. is brownish in colour, versus black in Unless otherwise stated explicitly, colour and other descriptions P. australis australis, or occasionally black with a very slight apply to living adult specimens of generally good health and not brown tinge only. under any form of stress by means such as excessive cool, Brown et al. (2006) at p. 312 describing P. australis heat, dehydration or abnormal skin reaction to chemical or other adelynhoserae sp. nov. as “The NQ gliders” separated the two input. subspecies as follows: Colour descriptions of species refer to fur colour (pelage) and “T he NQ glider s ar e smaller , as measur ed b y w eight (NQ not skin. Hoser 2020 - Australasian Journal of Herpetology 42:38-41. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 40 Australasian Journal of Herpetology

males: mean 516 g (s.d. = 7.7 g, n = 17); NQ females: Hoser, who glides like this subspecies of possum. She does this 479.4 g (s.d . = 7.8 g, n = 14) (Golding ay et al. 2001); V ic. as she does jumps on a snowboard at places like Whistler in males: 555.5 g (s.d . = 13.5 g , n = 11); Vic. f emales, 508.1 Canada, Hakuba in Japan and also the Australian ski resorts of g (s.d. = 8.1 g, n = 9 (M. Br own and S . M. Car thew, Thredbo, Mount Buller and Mount Hotham. unpublished da ta)), … There also a ppear to be REFERENCES CITED sociobeha viour al and life-history differ ences betw een the Anonymous 2008. Approved Conservation Advice (s266B of the NQ gliders and the southern forms. Environment Protection and Biodiversity Conservation Act 1999) T he NQ glider s ha ve been r epor ted to have a pol yg ynous Approved Conservation Advice for Petaurus australis unnamed m a ting system (Russell 1984; although see Golding ay et subsp. (Fluffy Glider). 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P., Wilson, D. E. and Reeder, D. M. (eds.) 2005. Mammal Species of the World: A Taxonomic and Geographic In terms of cranial characters, P. australis is separated from all Reference (3rd ed.). Johns Hopkins University Press, Baltimore, other species in the genus by having a large skull with; basal USA. length of more than 46 mm. Molars 1-3 are more than 7.3 mm long. Bullae are large, the posterior projecting lower than the Hedges, A. 2006. Biennial Yellow-Bellied Glider (Petaurus anterior part. australis reginae) Census, Daintree National Park: 2005 Results and an Overview of the Census since 1997. A report for the Thomas (1888) at pages 152-153 gives a more detailed Queensland Parks and Wildlife Service (QPWS), Mossman, description of P. australis, applicable to both subspecies. Queensland, Australia, March 2006. Distribution: Restricted to the wet tropics bioregion of Australia Hoser, R. T. 1991. Endangered Animals of Australia. Pierson from Mount Spec, north of Townsville, north Queensland in the Publishing, Mosman, NSW, Australia:240 pp. south, along the coastal rainforests, generally at high elevation running north to the Mount Windsor Tableland in the north. Hoser, R. T. 2019a. 11 new species, 4 new subspecies and a subgenus of Australian Dragon Lizard in the genus Etymology: Named in honour of my eldest daughter Adelyn

Tympanocryptis Peters, 1863, with a warning on the with sixty-five plates of non descript animals, birds, lizards, conservation status and long-term survival prospects of some serpents, curious cones of trees and other natural productions. newly named taxa. Australasian Journal of Herpetology 39:23- J. Debrett, London, UK. 52. Winter, J. W. 1997. Distribution of the Yellow-Bellied Glider Hoser, R. T. 2019b. Richard Shine et al. (1987), Hinrich Kaiser et Petaurus australis and Two Other Sympatric Petaurid Gliders in al. (2013), Jane Melville et al. (2018 and 2019): Australian the Wet Tropics Region of Queensland. A Report to the Wet Agamids and how rule breakers, liars, thieves, taxonomic Tropics Management Authority, April 1997. vandals and law breaking copyright infringers are causing reptile Winter, J. W. 2004, Forest mammals of northern Queensland: is species to become extinct. Australasian Journal of Herpetology their conservation status improving?, pp. 435-4451 in: Lunney, 39:53-63. D. (Ed.), Conservation of Australia’s Forest Fauna (second Iredale, T. and Troughton, E. L. G. 1934. A check-list of the edition): Royal Zoological Society of New South Wales, mammals recorded from Australia. Australian Museum Memoirs Mosman, Australia. 6:1-122. [4 May 1934]. Winter, J. W., Russell, R. A. W. and Winter, E. M. 1979. A Lawlor, T. E. 1979. Handbook to the Orders and Families of sighting of the yellow-bellied glider (Petaurus australis) in north- Living Mammals. Mad River Press, Inc. Eureka, California, USA. eastern Queensland. North Queensland Naturalist 45:2-4. Maxwell, S., Burbidge, A. A. and Morris, K. 1996. The 1996 Zaczek, Z. 2019. Former PM Kevin Rudd says Australia should Action Plan for Australian Marsupials and Monotremes. drastically increase its population to 50 MILLION to make sure Australasian Marsupial and Monotreme Specialist Group, IUCN nation can defend itself in the face of threats from China. Daily Species Survival Commission, Gland, Switzerland. Mail (Australia) (27 November), published online at: https:// Menkhorst, P. 2001. A Field Guide to the Mammals of Australia. www.dailymail.co.uk/news/article-7729689/Kevin-Rudd-says- Oxford University Press. Australia-increase-population-50-MILLION-defend-China.html Ride, W. D. L. 1970. A Guide to the Native Mammals of (last downloaded on 10 February 2020). Australia. Oxford University Press. CONFLICTS OF INTEREST Ride, W. D. L. (ed.) et al. (on behalf of the International None. Commission on Zoological Nomenclature) 1999. International code of Zoological Nomenclature. The Natural History Museum - Cromwell Road, London SW7 5BD, UK. Russell, R. 1980. Spotlight on possums. University of Queensland Press, Brisbane, Queensland, Australia. Saunders, S. 2019. Morrison and Shorten’s Big Australia: The overpopulation problem. News article published 17 April 2019 at: https://independentaustralia.net/politics/politics- display/morrison-and-shortens-big-australia-the- overpopulation-problem,12584 (last downloaded on 10 February 2020). Strahan, R. (ed.) 1988. The Australian Museum Complete book of Australian Mammals / The National Photographic Index of Australian Wildlife. Angus and Robertson, North Ryde, NSW, Australia: 530 pp. Tate, G. H. H. 1952. Mammals of Cape York Peninsula, with notes on the occurrence of rain forest in Queensland. Bulletin of the American Museum of Natural History 98:567-616. Thomas, O. 1888. Catalogue of the Marsupialia and Monotremata in the Collection of the British Museum (Natural History). British Museum, London, UK:xiii+401 pp+28 pl. Thomas, O. 1922. A selection of lectotypes of the typical Australian marsupials in the British Museum Collection. Ann. Mag. Nat. Hist. 9(10):127-128. Thomas, O. 1923. On some Queensland Phalangeridae. Ann. Mag. Nat. Hist. 9(11):246- 250. Vaughan, T. A. 1986. Mammalogy, 3rd Edition. Harcourt Brace Jovanovich College Publishers. New York, USA. White, J. 1790. Journal of a

Hoser 2020 - Australasian Journal of Herpetology 42:38-41. voyage to New South Wales : Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 42 Australasian Journal of Herpetology Australasian Journal of Herpetology 42:42-49. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

The ability to conserve a threatened species begins when they are named! New species of Rock Wallaby (Marsupialia: Macropodidae: Petrogale) from northern Australia.

LSID URN:LSID:ZOOBANK.ORG:PUB:24B992B7-1FF8-463E-8EF9-579A2C63DD78

RAYMOND T. HOSER

LSID urn:lsid:zoobank.org:author:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 20 February 2020, Accepted 16 March 2020, Published 25 April 2020.

ABSTRACT Numerous studies have been conducted into the phylogeny of Australia’s Rock Wallaby species within the widespread genus Petrogale Gray, 1837. In spite of these studies, well-known forms remain unnamed and therefore potentially under the radar of State and Federal wildlife conservation authorities, increasing their potential risk of decline or extinction. To rectify this situation, and in accordance with the rules of the International Code of Zoological Nomenclature, two highly divergent forms are formally named herein as species and two other populations as subspecies. Formally named for the first time are Petrogale martinekae sp. nov. from the west Kimberley in Western Australia, previously regarded as a southern population of the distinctive species known as the Warabi, P. burbidgei Kitchener and Sanson, 1978. Also formally named is Petrogale hoserae sp. nov. from the southern edge of the Gulf of Carpentaria, until now treated as a divergent population of the Eastern Short-eared Rock Wallaby P. wilkinsi Thomas, 1926. Two genetically and morphologically distinct east Kimberley populations of the Short-eared rock-wallaby P. brachyotis (Gould, 1841) are formally named as subspecies, being P. brachyotis pentecostensis subsp. nov. and P. brachyotis ordensis subsp. nov.. The formal naming of these taxa now enables wildlife departments to formulate conservation plans for extant populations and reduces the risk of their extinctions arising from them being treated as one and the same as otherwise more widespread and abundant species. This paper also formally divides the genus into four subgenera based on known divergences of each major species group. Two subgenera are formally named for the first time. Keywords: Taxonomy; nomenclature; classification; Wallaby; Marsupials; Rock Wallaby; northern Australia; Australia; Western Australia; Northern Territory; Queensland; Macropodidae; Petrogale; burbidgei; brachyotis; wilkinsi; persephone; xanthopus; celeris; new subgenus; Quasipetrogale; Ferepetrogale; new species; hoserae; martinekae; new subspecies; pentecostensis; ordensis.

INTRODUCTION increasing their potential risk of decline or extinction. Rock Wallabies of the genus Petrogale Gray, 1837 are To rectify this situation, and in accordance with the rules of the widespread and common in most parts of Australia. International Code of Zoological Nomenclature, the genus was Numerous studies have been conducted into the phylogeny of reviewed and as a result two highly divergent forms were Petrogale Gray, 1837, including those cited in the materials and identified as potentially unnamed species as were a number of methods section of this paper. populations divergent at the subspecies level. In spite of these studies, well-known and previously identified As this paper was being prepared, some of these forms were in divergent forms remain unnamed and therefore potentially under fact formally named (Eldridge and Potter, 2020), which the radar of State and Federal wildlife conservation authorities, necessitated removal of two subspecies descriptions from this

paper in the pre-publication process. Live and dead specimens as well as available bone specimens, Remaining were at least two other putative species and two were examined as was other necessary material, including past potential subspecies. climate data for the relevant regions, sea level depths, and other The relevant candidate taxa were examined in terms of their relevant information. existing classification and also in terms of the published In summary, as inferred already, the genetic, geological, literature, phylogenies, divergences and available names in historical and morphological evidence clearly showed that there terms of the International Code of Zoological Nomenclature were two obviously unnamed species of Rock Wallaby in the P. (Ride et al. 1999), before the final decision was made to publish brachyotis (Gould, 1841) species group. this paper. The south western Kimberley Ranges population of the Formally identified as new taxa worthy of being named and distinctive species known as the Warabi, P. burbidgei Kitchener named for the first time are Petrogale martinekae sp. nov. from and Sanson, 1978 was sufficiently divergent from the type form the west Kimberley in Western Australia, formerly regarded as a from the Mitchell Plateau to warrant being recognized as a full southern population of the distinctive species known as the species. It is herein formally named Petrogale martinekae sp. Warabi, P. burbidgei Kitchener and Sanson, 1978. Also formally nov. with a known distribution from the Prince Regent River in named is Petrogale hoserae sp. nov. from the southern edge of the north to the Munboon Plateau in the south (the type locality). the Gulf of Carpentaria, until now treated as a divergent A very divergent population of the Eastern Short-eared Rock population of the Eastern Short-eared Rock Wallaby P. wilkinsi Wallaby P. wilkinsi Thomas, 1926 from the southern shores of Thomas, 1926. the Gulf of Carpentaria was also found to be sufficiently Two genetically and morphologically distinct east Kimberley divergent from that taxon with a type locality of the top end of the populations of Short-eared rock-wallaby P. brachyotis (Gould, Northern Territory, to warrant being identified and named as a 1841) are formally named as subspecies, being P. brachyotis new species. pentecostensis subsp. nov. and P. brachyotis ordensis subsp. It is formally named as Petrogale hoserae sp. nov.. nov.. Two genetically and morphologically distinct east Kimberley The formal naming of these species now enables wildlife populations of Short-eared rock-wallaby P. brachyotis (Gould, departments to formulate conservation plans for extant 1841) as identified by Potter et al. (2014) are herein formally populations of these taxa and reduces the risk of their named as subspecies. extinctions arising from them being treated as one and the same as otherwise more widespread and abundant species. These are P. brachyotis pentecostensis subsp. nov. and P. brachyotis ordensis subsp. nov. named in reflection of the At the genus level, Petrogale has long been known to consist of known centres of distribution for each subspecies. three (according to most authors) or four (according to Potter et Potter et al. (2014) identified each population as EK1 ESU and al. 2012) main species groups. EK2 ESU. However use of subgeneric classification has been rarely used Both are potentially threatened due to their relatively limited for these groups within Petrogale. known distributions and ongoing risk of decline from introduced This remains the case, even though Potter et al. (2012), showed species such as foxes, cats (in particular) (see Spencer 1991), subgenus-level divergences for four main clusters of species. dogs or direct human intervention in other ways. The name Peradorcas Thomas, 1904 with the type species P. Significantly, the four newly named taxa have all diverged across concinna Gould, 1842, is available for the so-called P. brachyotis known biogeographical barriers that have resulted in numerous (Gould, 1841) species group. No names are available for the species of reptile being discovered and formally named in recent other two divergent lineages and they are herein formally named years, having been separated from their nearest relatives across for the first time. exactly the same barriers. MATERIALS, METHODS AND RESULTS Examples can be seen in the new species descriptions within Before a decision is made to name any new taxon, reasonable Hoser (2014, 2015, 2016, 2017, 2018a, 2018b, 2018c, 2018d). steps must be taken to ensure that it is justified on all relevant As mentioned in the abstract the formal naming of these species grounds, including that it is morphologically, genetically and now enables wildlife departments to formulate conservation reproductively isolated from their nearest relative and to a plans for extant populations of these taxa and reduces the risk sufficient degree to be of taxonomic significance. of their extinctions arising from them being treated as one and A further relevant question to ask is should the reproductively the same as otherwise more widespread and abundant species isolated and morphologically divergent entities be labelled as as has previously occurred (see Hoser 2019a, 2019b). subspecies, full species, or potentially higher level again. The genus Petrogale has long been recognized as consisting Key literature relevant to the taxonomic and nomenclatural four morphologically distinct groups potentially worthy of conclusions within this paper include Bannister et al. (1988), subgeneric ranking or potentially even genus-level division Bee and Close (1993), Browning et al. (2001), Bryant and (Thomas 1909). Perhaps due to the presence of the pre-existing Krosch (2016), Burbidge and Manly (2002), Burbidge and synonym for the second main group of species, no recent McKenzie (1978, 1989), Collins (1973), Clancy and Close publishing authors since Thomas (1904) have utilized subgenus- (1997), Eldridge (1997), Eldridge and Close (1992, 1993, 1997), level classifications for the genus. Eldridge and Potter (2020), Eldridge et al. (1992a, 1992b, 2001, The P. penicillata (Gray, 1825) group is the type group for the 2012), Flannery and Archer (1984), Flannery et al. (1992), genus and so would be the nominate subgenus if the genus Gibson and McKenzie (2012), Goodfellow (1993), Gould (1841, were to be divided. 1842), Gray (1837), Groves et al. (2005), Hoser (1991), Iredale and Troughton (1934), Kitchener and Sanson (1978), Lawlor The name Peradorcas Thomas, 1904 is available for the P. (1979), Le Souef (1924), Maynes (1982), Maxwell et al. (1996), concinna Gould, 1842 species group, better-known to most Menkhorst (2001), Meredith et al. (2008), Pearson (2012), Potter people as the P. brachyotis (Gould, 1841) species group (see et al. (2012a, 2012b, 2012c, 2014), Ride (1970), Ride et al. below). (1999), Roache (2011), Sharman et al. (1990), Shuker and The molecular phylogeny published by Potter et al. (2012) at Durrell (1993), Spencer (1991), Start et al. (2012), Strahan Fig. 5 showed subgenus level divergences within Petrogale for (1988), Telfer and Eldridge (2010), Thomas (1888, 1904a, four species groups. These were for the two groups mentioned 1904b, 1909, 1922, 1926a, 1926b), Vaughan (1986) and above as well as two other divergent lineages for two other sources cited therein (duplicitous references not necessarily divergent species, namely P. xanthopus Gray, 1855 and P. included). persophone Maynes, 1982. Potter et al. (2012) wrote: “Four distinct lineages were identified, Hoser 2020 - Australasian Journal of Herpetology 42:42-49. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 44 Australasian Journal of Herpetology

(1) the brachyotis group, (2) Petrogale persephone, (3) The tail also terminates in a white to yellowish-white tip 15-20 Petrogale xanthopus and (4) the lateralis-penicillata group.” mm in length (in adults). Head and body length (in adults) is 52- Therefore this paper also formally divides the genus Petrogale 64 cm, tail length (in adults) 60-68 cm and weight (in adults) is into four subgenera based on known divergences of each major 5-8 kg with males on average 60% heavier than females species group. Two subgenera are formally named for the first (adapted from Strahan 1988). time, namely Quazipetrogale subgen. nov. and Ferepetrogale The genus Petrogale is diagnosed and defined in Thomas subgen. nov.. (1888) at pages 62-64. Potter et al. (2012) showed a divergence of about 8 MYA from Distribution: Living specimens are known only from a small other species for each of these subgenus-level groups, which area in Conway National Park, Dryander National Park, was broadly in line with the marginally more divergent subgenus Gloucester Island National Park and around the town of Airlie Peradorcas. Beach, all in Whitsunday Shire in Queensland, Australia. INFORMATION RELEVANT TO THE FORMAL DESCRIPTIONS Etymology: Quasi means “apparently but not really; seemingly.” THAT FOLLOW which fits in line with the similarity of this species and subgenus There is no conflict of interest in terms of this paper or the to the type species in the genus Petrogale Gray, 1837. conclusions arrived at herein. Content: Petrogale (Quasipetrogale) persephone Maynes, 1982 Several people including anonymous peer reviewers who (monotypic). revised the manuscript prior to publication are also thanked as FEREPETROGALE SUBGEN. NOV. are relevant staff at museums who made specimens and LSID urn:lsid:zoobank.org:act:D1643CB0-3FD6-415F-89CD- records available in line with international obligations. A5F1129B7838 In terms of the following formal descriptions, spellings should Type Species: Petrogale xanthopus Gray, 1855. not be altered in any way for any purpose unless expressly and exclusively called for by the rules governing Zoological Diagnosis: The two morphologically similar species within the Nomenclature as administered by the International Commission subgenus Ferepetrogale subgen. nov. are readily separated of Zoological Nomenclature. from all other species (and subgenera within the genus Petrogale Gray, 1837) by the following unique combination of In the unlikely event two or more newly named taxa are deemed characters: Tail is annulated with brown and pale yellow being conspecific by a first reviser, then the name to be used and the only two species in the genus Petrogale with this character. retained is that which first appears in this paper by way of page There is a dark nuchal stripe present. Ears are uniform yellow priority and as listed in the abstract keywords. from behind. Skull length (in adults) is more than 9 cm long, the Some material in descriptions for taxa may be repeated for other muzzle is broad, its sides and the interorbital region inflated. taxa in this paper and this is necessary to ensure each fully Greatest breadth of nasals is about one third of their length. complies with the provisions of the International Code of There is a rich brown mid-dorsal stripe running from the crown of Zoological Nomenclature (Fourth edition) (Ride et al. 1999) as the head to the centre of the back, a distinct white cheek mark; amended online since. brick-red patch running narrowly from the upper arm and onto Material downloaded from the internet and cited anywhere in this the elbows and forearms, at which stage it effectively encircles paper as being sourced online was downloaded and checked the limb and there is also a buffy white side stripe followed by a most recently as of 10 March 2020 (including if also viewed brown hip stripe (modified from Thomas 1888, Strahan 1988). prior), unless otherwise stated and was accurate in terms of the Distribution: Found around the Southern Gulfs of South content (as described) cited herein as of that date. Australia, Bulloo River basin and the Murray/Darling river basin Unless otherwise stated explicitly, colour and other descriptions in New South Wales and Queensland, Australia. apply to living adult specimens of generally good health and not Etymology: Fere, in Latin means not quite, or almost, hence the under any form of stress by means such as excessive cool, name Ferepetrogale literally means not quite Petrogale. heat, dehydration or abnormal skin reaction to chemical or other Content: Petrogale (Ferepetrogale) xanthopus Gray, 1855 (type input. Colour descriptions of species refer to fur, not skin. species); P. (Ferepetrogale) celeris Le Souef, 1924. While numerous texts and references were consulted prior to PETROGALE (PERADORCAS) HOSERAE SP. NOV. publication of this paper, the criteria used to separate the relevant species or subspecies has already been spelt out and/ LSID urn:lsid:zoobank.org:act:BD1569F2-126A-433F-B912- or is done so within each formal description and does not rely on 3FCEE388F73E material within publications not explicitly cited herein. Holotype: A preserved specimen at the Australian Museum in QUASIPETROGALE SUBGEN. NOV. Sydney, New South Wales, Australia, specimen number M.38966 collected at Moonlight Gorge, Wollogorang, Northern LSID urn:lsid:zoobank.org:act:7780E296-D22F-4D39-83E3- Territory, Australia, Latitude -17.20 S., Longitude 137.80 E. This 73E8CC8C9B25 government-owned facility allows access to its holdings. Type Species: Petrogale persephone Maynes, 1982. Paratype: A preserved female specimen at the Australian Diagnosis: The single species within the subgenus Museum in Sydney, New South Wales, Australia, specimen Quasipetrogale subgen. nov. is readily separated from all other number M.45290.001 collected at Red Bank Mine, Wollogorang, species (and subgenera within the genus Petrogale Gray, 1837) N.T., Australia, Latitude -17.18 S., Longitude 137.73 E. by the following unique combination of characters: ears with Diagnosis: The species Petrogale hoserae sp. nov. has until brownish-orange hair outside and black internally. White to pale now been treated as a number of other taxa, most notably P. yellowish-white or pale whitish-grey lateral stripe on upper lip, brachyotis (Gould, 1841) until the review of the species group by passing below eye to about level of ear with platinum or pearl Potter et al. (2014). grey below which arises at the corner of the mouth and passes back to the ear region. Shoulders raw sienna passing to dark While finding the lineage herein described as a new species to brown in axillary region with rest of the dorsal surface light be divergent from the species P. wilkinsi Thomas, 1926 (from brown, lightly pencilled with white hairs. Chin is white and throat the top end of the Northern Territory), Potter et al. (2014) placed varies from white to off-white, chest and venter light yellow to this taxon and another described form, namely P. longmani creamy yellow. Toes of fore and hind limbs and about half of Thomas, 1926 from Groote Eylandt, Northern into the hind feet are black. Recently moulted specimens may be greyish synonymy of P. wilkinsi. rather than brownish in overall pelage. The species presents as On the basis of morphological, molecular and reproductive a generally dark animal, made such by the overall colour divergence, all three of these forms, all occurring in the Northern combined with the black feet and black dorsal surface of the tail. Territory are herein treated as separate species. Hoser 2020 - Australasian Journal of Herpetology 42:42-49. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 45

All three species fit within the diagnosis of P. wilkinsi as Distribution: P. hoserae sp. nov. occurs from near Hells Gate, published within Potter et al. (2014). near Nicholson, far north-west Queensland, west along the The three species are readily separated from one another and southern edge of the Gulf of Carpentaria, through the type other species in the subgenus Peradorcas on the basis of locality at Wollogorang, Northern Territory, where suitable hilly colouration which is as follows: and rocky habitat occurs, to about Ngukurr, Walmudga Hill, P. wilkinsi, P. longmani and P. hoserae sp. nov. are all readily 10km east of Ngukurr (Roper River Mission) and the nearby Sir separated from both P. victoriae (Eldridge and Potter, 2014) Edward Pellew Islands. (treated here as a full species as opposed a subspecies of P. Etymology: Named in honour of my mother, Katrina Joan brachyotis as originally named) and all subspecies of P. Hoser, of Lane Cove, New South Wales, Australia, for services brachyotis by the following suite of characters: to wildlife conservation spanning more than 50 years. Dorsal surface is dark grey to grey/brown (P. wilkinsi), or PETROGALE (PERADORCAS) MARTINEKAE SP. NOV. silverish-grey in colour (P. longmani), or a yellowish-brown colour LSID urn:lsid:zoobank.org:act:4E88DBDF-B805-401A-B767- (P. hoserae sp. nov.). In P. wilkinsi and P. longmani the ventral 65B1751D1A6A surfaces are distinctly marked, while in P. hoserae sp. nov. they Holotype: A preserved specimen at the Australian Museum in are less so. In P. wilkinsi and P. longmani the dorsal stripe is a Sydney, New South Wales, Australia, specimen number prominent dark brown/black colour, running from forehead to M.47136.001, collected from the edge of the Munboon Plateau, beyond shoulders and slightly less so in P. hoserae sp. nov.. Western Australia, Latitude -16.4 S., Longitude 125.1 E. This Likewise for the distinct white shoulder stripe behind a well government-owned facility allows access to its holdings. developed dark axillary patch, which is present, but slightly less Paratype: A preserved specimen at the Western Australian distinct in P. hoserae sp. nov.. Museum in Perth, Western Australia, Australia, specimen Limbs are distinctly and often brightly coloured; from yellow number M54836 collected from the Prince Regent River -15.6 through orange to reddish brown. Face is distinct, being light S., Longitude 125.3 E. brown to orangey brown in P. wilkinsi and P. longmani, while Diagnosis: Petrogale martinekae sp. nov. has until now been being a dark brown to blackish in P. hoserae sp. nov.. treated as a southern population of a lineage of P. burbidgei The tail is lighter than the body on the dorsal surface, sides Kitchener and Sanson, 1978, as defined by those authors. more yellowish. Terminal 1/5 to 1/3 tail is darker; brown to However Potter et al. (2014) identified this taxon as being almost black, especially towards the tip. divergent enough to warrant species-level identification and for P. longmani is separated from both P. wilkinsi and P. hoserae sp. which no name was available (Bannister et al. 1988). nov. by its overall silvery-grey colour, including a grey shoulder P. martinekae sp. nov. is similar in most respects to P. burbidgei stripe and axillary patch that is black; dorsal stripe being thick and would until now be identified as that taxon. P. martinekae sp. and dark, extending to the mid back and enlarged into an nov.. It is however separated from P. burbidgei by the following irregular dark patch on top of the head. suite of adult characters in life: the inner ears have a distinct P. hoserae sp. nov. is separated from both P. wilkinsi and P. light bluish tinge, versus pink in P. burbidgei; while both species longmani by its distinctively light brown colour dorsally (or are brownish in pelage, the outer grey to black peppering is alternatively yellow), sometimes with a strong reddish tinge, prominent in P. martinekae sp. nov. and indistinct in P. burbidgei. markings, including ventral ones, are clearly faded and not The upper hind feet are greyish black in P. martinekae sp. nov. prominent and the tail and face are also exceptionally dark. versus dark brown in P. burbidgei. The fur of the centre of the P. brachyotis (all subspecies) and P. victoriae (Eldridge and chest in P. martinekae sp. nov. is white, versus off-white in P. Potter, 2014) (treated here as a full species as opposed a burbidgei. The black line running from the nose to the crown is subspecies of P. brachyotis as originally named) are separated well-defined in P. martinekae sp. nov. versus ill-defined in P. from all other species in the subgenus Peradorcas by the burbidgei. following characters: Back of ears uniform grey or fawn, P. martinekae sp. nov. and P. burbidgei are morphologically sometimes tipped with white. No occipital or nuchal streak. similar to P. concinna Gould, 1842, but both are readily Shoulder markings are present. separated from that taxon (all subspecies) by having obviously Frontal outline is convex or straight above the front of the orbit. shorter ears, being less than 35 mm long, versus greater than Nasals are expanded behind. 35 mm in P. concinna. Type P. brachyotis (Gould, 1841) have a P. victoriae are readily separated from P. brachyotis (all well-defined dark brown neck stripe to about the level of the subspecies) by the following suite of characters: yellowish-grey shoulder, as does the east Kimberley lineage herein named P. colour dorsally; no side or hip stripes; virtually no ventral brachyotis pentecostensis subsp. nov.. This is either indistinct or markings; limbs are same colour as body, or slightly more only prominent on the elbow region in P. martinekae sp. nov. and yellowish; face is slightly darker than neck and shoulders; the P. burbidgei. The taxon P. brachyotis ordensis subsp. nov. is tail is the same colour as the body or lighter, then becoming a separated from P. martinekae sp. nov. and P. burbidgei by darker brown at the terminal third to fifth. having well-defined face markings versus indistinct in P. martinekae sp. nov. and P. burbidgei. P. martinekae sp. nov. (until now treated as a population of P. burbidgei), P. burbidgei and P. concinna are separated from P. P. martinekae sp. nov., P. burbidgei and P. concinna are brachyotis (all subspecies), P. victoriae (Eldridge and Potter, separated from P. brachyotis (all subspecies), P. victoriae 2014) (treated here as a full species as opposed a subspecies (Eldridge and Potter, 2014) (treated here as a full species as of P. brachyotis as originally named), P. wilkinsi Thomas, 1926, opposed a subspecies of P. brachyotis as originally named), P. P. longmani Thomas, 1926 and P. hoserae sp. nov. by their wilkinsi Thomas, 1926, P. longmani Thomas, 1926 and P. smaller size, the hind foot being 105 mm long or smaller, versus hoserae sp. nov. by their smaller size, the hind foot being 105 over 105 mm in the other species. mm long or smaller, versus over 105 mm in the other species. P. martinekae sp. nov. can be seen in life in images at: P. victoriae (Eldridge and Potter, 2014) (treated here as a full species as opposed a subspecies of P. brachyotis as originally https://www.flickr.com/photos/12742129@N07/49114207146/ named) and P. brachyotis are separated from all of P. hoserae and sp. nov., P. wilkinsi and P. longmani by having no side stripe or https://www.flickr.com/photos/12742129@N07/49114420862/ hip stripe, versus a pale one in P. hoserae sp. nov., P. wilkinsi and and P. longmani, a grey (instead of white) shoulder stripe; an indistinct dorsal stripe (versus obvious) and a greyish white https://www.flickr.com/photos/12742129@N07/49114387852/ venter (instead of white). P. burbidgei Kitchener and Sanson, 1978 can be seen in life in Hoser 2020 - Australasian Journal of Herpetology 42:42-49. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 46 Australasian Journal of Herpetology images at: Potter, 2014) (treated here as a full species as opposed a https://www.flickr.com/photos/126002448@N02/21199638021/ subspecies of P. brachyotis as originally named) are separated and from all other species in the subgenus Peradorcas by the https://www.flickr.com/photos/88708273@N03/48945453017/ following characters: Back of ears uniform grey or fawn, sometimes tipped with white. No occipital or nuchal streak. and Shoulder markings are present. https://www.flickr.com/photos/reptileshots/14533826956/ Frontal outline is convex or straight above the front of the orbit. and Nasals are expanded behind, https://www.flickr.com/photos/reptileshots/14584554932/ P. victoriae are readily separated from P. brachyotis (all and subspecies) by the following suite of characters: yellowish-grey https://www.flickr.com/photos/88708273@N03/48944717823/ colour dorsally; no side or hip stripes; virtually no ventral markings; limbs are same colour as body, or slightly more Distribution: P. martinekae sp. nov. is known only from the yellowish; face is slightly darker than neck and shoulders; the region running from the Prince Regent River to the Munboon tail is the same colour as the body or lighter, then becoming a Plateau in the west Kimberley district of Western Australia. darker brown at the terminal third to fifth. P. burbidgei Kitchener and Sanson, 1978 is found from the Images of nominate Petrogale brachyotis brachyotis in life can Prince Regent River north to the Mitchell Plateau, including be found online at: some offshore islands. https://www.flickr.com/photos/123952930@N04/44125908541/ Etymology: Named in honour of Maryann Martinek, in 2020 of Bendigo, Australia, in recognition of her services to wildlife and conservation globally. For more detail refer to Hoser (2010). https://www.flickr.com/photos/ailognom/39807952372/ PETROGALE (PERADORCAS) BRACHYOTIS and PENTECOSTENSIS SUBSP. NOV. https://www.flickr.com/photos/ailognom/39807959912/ LSID urn:lsid:zoobank.org:act:2DB42ACD-F0E3-453F-AFD4- An image of P. brachyotis pentecostensis subsp. nov. in life can ACC526A97859 be seen at: Holotype: A preserved female specimen at the Australian https://splashingpaint.files.wordpress.com/2019/06/ Museum in Sydney, New South Wales, Australia, specimen p_wa9335x.jpg number M.45275.003 collected at El Questro Station, Pentecost Images of P. brachyotis ordensis subsp. nov. in life can be seen River, Kimberley District, Western Australia, Australia, Latitude - at: 16.03 S., Longitude 127.98 S. https://www.flickr.com/photos/58349528@N02/44492360964/ This government owned facility allows access to its holdings. and Paratypes: Five preserved specimens at the Australian Museum https://www.flickr.com/photos/58349528@N02/45163729682/ in Sydney, New South Wales, Australia, specimen numbers M.45274.004, M.45272.003, M.45271.002, M.45273.002 and and M.45274.004 collected from at or immediately adjacent to (within https://www.flickr.com/photos/58349528@N02/44302861675/ 10 km) the type locality given above. Distribution: P. brachyotis pentecostensis subsp. nov. is known Diagnosis: Until now the subspecies Petrogale brachyotis only from the immediate vicinity of the type locality as well as the pentecostensis subsp. nov. has been treated as an easterly Ord basin (known from Monsmont Island in Lake Argyle) and population of nominate P. brachyotis (Gould, 1841), although I assumed to occupy only this general region and slightly west of note Eldridge and Potter (2014) found subspecies level mtDNA there to the Durack River. divergence for this population as compared to another Etymology: Named after the type locality, the Pentecost River. population east of this one, being east of the Ord River and PETROGALE (PERADORCAS) BRACHYOTIS ORDENSIS herein named as P. brachyotis ordensis subsp. nov. and the SUBSP. NOV. nominate P. brachyotis from the north-west Kimberley district. LSID urn:lsid:zoobank.org:act:64577A73-8436-493E-85D3- All three can be readily separated on the basis of colouration. 40D87F1FD1B6 Nominate P. brachyotis brachyotis is separated from P. Holotype: A preserved specimen at the Western Australian brachyotis pentecostensis subsp. nov. and P. brachyotis Museum Mammal Collection, Perth, Western Australia, ordensis subsp. nov. by the following characters: having a Australia, specimen number M19865 collected from Point reddish-brown dorsal colour, very well defined side markings, Spring, Western Australia, Australia, Latitude -15.41 S., distinctively orangeish hind limbs, generally dark greyish head Longitude 128.89 E. This government owned facility allows and orangish grey inner ears. access to its holdings. P. brachyotis pentecostensis subsp. nov. is separated from P. Paratype: A preserved specimen at the Western Australian brachyotis brachyotis and P. brachyotis ordensis subsp. nov. by Museum Mammal Collection, Perth, Western Australia, the following characters: having a greyish dorsum with limited Australia, specimen number M24401 collected from Ninbing amounts of orange or russet dusting on the hind limbs, Bore, WA, Australia, Latitude -15.41 S., Longitude 128.40 E. moderately well defined side markings, a face that is no darker than the body with ill defined and light, darker markings on the Diagnosis: Until now the subspecies Petrogale brachyotis snout (versus dark and well defined in the other two subspecies) pentecostensis subsp. nov. has been treated as an easterly and a brownish head and brown inner ears. population of nominate P. brachyotis (Gould, 1841), although I note Eldridge and Potter (2014) found subspecies level mtDNA P. brachyotis ordensis subsp. nov. is separated from P. divergence for this population as compared to another brachyotis pentecostensis subsp. nov. and P. brachyotis population east of this one, being east of the Ord River and brachyotis by the following characters: having a light greyish herein named as P. brachyotis ordensis subsp. nov. and the dorsum, flecked with either yellow or orangeish, with brown, nominate P. brachyotis from the north-west Kimberley district. rather than orange on the hind limbs, ill defined side markings (versus well defined in P. brachyotis pentecostensis subsp. nov. All three can be readily separated on the basis of colouration. and P. brachyotis), a generally greyish-beige head, being dark at Nominate P. brachyotis brachyotis is separated from P. tip of snout and between snout and eye and lighter elsewhere brachyotis pentecostensis subsp. nov. and P. brachyotis and a greyish-purple inner ear. ordensis subsp. nov. by the following characters: having a P. brachyotis (all subspecies) and P. victoriae (Eldridge and reddish-brown dorsal colour, very well defined side markings, Hoser 2020 - Australasian Journal of Herpetology 42:42-49. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 47

distinctively orangeish hind limbs, generally dark greyish head world of 2020. and orangish grey inner ears. Wildlife laws as currently enforced in Australia are not in any P. brachyotis pentecostensis subsp. nov. is separated from P. materially significant way enhancing the long-term survival brachyotis brachyotis and P. brachyotis ordensis subsp. nov. by prospects of any of the relevant species and in any event are the following characters: having a greyish dorsum with limited being vastly outweighed by other negative impacts of amounts of orange or russet dusting on the hind limbs, governments, including their ongoing commitment to growing moderately well defined side markings, a face that is no darker the human population to a level that can only put further than the body with ill defined and light, darker markings on the pressure on the survival prospects of the relevant taxa snout (versus dark and well defined in the other two subspecies) If the Australian government persists with its “Big Australia and a brownish head and brown inner ears. Policy”, (see for example Saunders 2019 or Zaczek 2019), that P. brachyotis ordensis subsp. nov. is separated from P. being a long-term aim to increase the human population in brachyotis pentecostensis subsp. nov. and P. brachyotis Australia to over 100 million people by year 2150 (from the brachyotis by the following characters: having a light greyish present 25 million as of 2019), all sorts of unforseen threats to dorsum, flecked with either yellow or orangeish, with brown, the survival of these species will almost certainly emerge. rather than orange on the hind limbs, ill defined side markings REFERENCES CITED (versus well defined in P. brachyotis pentecostensis subsp. nov. Bannister, J. L., Calaby, J. H., Dawson, L. J., Ling, J. K., and P. brachyotis), a generally greyish-beige head, being dark at Mahoney, J. A., McKay, G. M., Richardson, B. J., Ride, W. D. L. tip of snout and between snout and eye and lighter elsewhere and Walton, D. M. 1988. Zoological Catalogue of Australia, and a greyish-purple inner ear. Volume 5, Mammals. Australian Government Publishing Service, P. brachyotis (all subspecies) and P. victoriae (Eldridge and Canberra, ACT, Australia:X+274 pp. Potter, 2014) (treated here as a full species as opposed a Bee, C. A. and Close, R. L. 1993. Mitochondrial DNA analysis of subspecies of P. brachyotis as originally named) are separated introgression between adjacent taxa of rock-wallabies, Petrogale from all other species in the subgenus Peradorcas by the species. Genetical Research 61:21-37. following characters: Back of ears uniform grey or fawn, Browning, T. L., Taggart, D. A., Rummery, C., Close, R. L. and sometimes tipped with white. No occipital or nuchal streak. Eldridge, M. D. B. 2001. Multifaceted genetic analysis of the Shoulder markings are present. critically endangered brush-tailed rock-wallaby Petrogale Frontal outline is convex or straight above the front of the orbit. penicillata in Victoria, Australia: implications for management. Nasals are expanded behind. Conservation Genetics 2:145-156. P. victoriae are readily separated from P. brachyotis (all Bryant, L. M. and Krosch, M. N. 2016. Lines in the land: a review subspecies) by the following suite of characters: yellowish-grey of evidence for eastern Australia’s major biogeographical colour dorsally; no side or hip stripes; virtually no ventral barriers to closed forest taxa. Biological Journal of the Linnean markings; limbs are same colour as body, or slightly more Society, 2016:27 pp. (online, downloaded most recently on 10 yellowish; face is slightly darker than neck and shoulders; the March 2020). tail is the same colour as the body or lighter, then becoming a Burbidge, A. A. and Manly, B. F.J. 2002. Mammal extinctions on darker brown at the terminal third to fifth. Australian islands: causes and conservation implications. Images of nominate Petrogale brachyotis brachyotis in life can Journal of Biogeography 29:465-474. be found online at: Burbidge, A. A. and McKenzie, N. 1978. The islands of the https://www.flickr.com/photos/123952930@N04/44125908541/ North-west Kimberley Western Australia. Wildlife Research and Bulletin of Western Australia 7:1-47. https://www.flickr.com/photos/ailognom/39807952372/ Burbidge, A. A. and McKenzie, N. L. 1989. Patterns in the and modern decline of Western Australia’s vertebrate fauna: causes https://www.flickr.com/photos/ailognom/39807959912/ and conservation implications. Biological Conservation 50:143- 198. An image of P. brachyotis pentecostensis subsp. nov. in life can Collins, L. R. 1973. Monotremes and Marsupials. Smithsonian be seen at: Institution Press. City of Washington, DC, USA. https://splashingpaint.files.wordpress.com/2019/06/ p_wa9335x.jpg Clancy, T. F. and Close, R. L. 1997. The Queensland Rock- wallabies - An overview of their conservation status, threats and Images of P. brachyotis ordensis subsp. nov. in life can be seen management. Australian Mammalogy 19:169-173. at: Eldridge, M. D. B. 1997. Taxonomy of rock wallabies Petrogale https://www.flickr.com/photos/58349528@N02/44492360964/ (Marsupialia: Macropodidae). II. An historical review. Australian and Mammalogy 19:113-122. https://www.flickr.com/photos/58349528@N02/45163729682/ Eldridge, M. D. B. and Close, R. L. 1992. Taxonomy of rock and wallabies, Petrogale (Marsupialia: Macropodidae). I. Revision of https://www.flickr.com/photos/58349528@N02/44302861675/ the eastern Petrogale with the description of three new species. Australian Journal of Zoology 40:605-624. Distribution: Petrogale brachyotis ordensis subsp. nov. is known only from the region between the lower Ord River, Eldridge, M. D. B. and Close, R. L. 1993. Radiation of Western Australia and the Bullo River in the Northern Territory. chromosome shuffles. Current Opinion in Genetics and Development 3:915-922. Etymology: Named in reflection of the region the subspecies occurs and is most commonly seen, being the lower Ord River Eldridge, M. D. B. and Close, R. L. 1997. Chromosomes and area in Western Australia, as in near Kununurra. evolution in rock-wallabies, Petrogale (Marsupialia: Macropodidae). Australian Mammalogy 19:123-135. CONSERVATION STATUS AND LONG TERM SURVIVAL PROSPECTS OF EACH DESCRIBED TAXON ABOVE Eldridge, M. D. B. and Potter, S. 2020. Taxonomy of rock- wallabies, Petrogale (Marsupialia: Macropodidae). V. A In terms of conservation of each population of each species or description of two new subspecies of the black-footed rock- subspecies as described above, the comments in Hoser (1991, wallaby (Petrogale lateralis). Australian Journal of Zoology 1993, 1996, 2019a and 2019b) apply. 67(1):19-26. None of the above taxa are regarded as under immediate risk of Eldridge, M. D. B., Johnston, P. G. and Close, R. L. 1992a. extinction, as best as such a judgement can be made in the

Chromosomal rearrangements in rock wallabies Petrogale Hoser, R. T. 2016. Carphodactylidae reviewed: Four new genera, (Marsupialia: Macropodidae) VI. Determination of the four new subgenera, nine new species and four new subspecies plesiomorphic karyotype for Petrogale. G-banding comparison of within the Australian gecko family (Squamata: Sauria). Thylogale with P. persephone, P. xanthopus and P. l. lateralis. Australasian Journal of Herpetology 32:3-25. Cytogenetics and Cell Genetics 61:29-33. Hoser, R. T. 2017. A further break-up of the Australian gecko Eldridge, M. D. B., Johnston, P. G. and Lowry, P. S. 1992b. genus Oedura Gray, 1842 Chromosomal rearrangements in rock wallabies, Petrogale sensu lato as currently recognized, from four to seven genera, (Marsupialia: Macropodidae). VII. G-banding analysis of with two new subgenera defined, description of fourteen new Petrogale brachyotis and P. concinna: species with dramatically species, four new subspecies and formalising of one tribe and altered karyotypes. Cytogenetics and Cell Genetics 61:34-39. five subtribes. Australasian Journal of Herpetology 34:3-35. Eldridge, M. D. B., Wilson, A. C. C., Metcalfe, C. J., Dollin, A. Hoser, R. T. 2018a. Six new species of Dwarf Goanna, E., Bell, J. N., Johnson, P. M., Worrellisaurus Wells and Wellington, 1984 from Australia. Johnston, P. G. and Close, R. L. 2001. Taxonomy of rock- Australasian Journal of Herpetology 37:24-37. wallabies, Petrogale (Marsupialia: Macropodidae). III. Molecular Hoser, R. T. 2018b. A new species within the Odatria glauerti data confirms the species status of the purple-necked rock- (Squamata: Varanidae) species complex. Australasian Journal wallaby (Petrogale purpureicollis Le Souef). Australian Journal of of Herpetology 37:44-45. Zoology 49:323-343. Hoser, R. T. 2018c. The description of a fourth subspecies of Eldridge, M. D. B., Potter, S. and Cooper, S. J. B. 2012. Odatria (Kimberleyvaranus) glebopalma (Mitchell, 1955) Biogeographic barriers in north-western Australia: an overview (Reptilia: Squamata: Varanidae). Australasian Journal of and standardization of nomenclature. Australian Journal of Herpetology 37:46-47. Zoology 59:270-272. Hoser, R. T. 2018d. A divided Gehyra makes sense! Assigning Flannery, T. F. and Archer, M. 1984. The macropodoids available and new names to recognize all major species groups (Marsupialia) of the early Pliocene Bow Local Fauna, central within Gehyra Gray, 1834 sensu lato (Squamata: Gekkonidae) eastern New South Wales. Aust. Zool. 21:357-383. and the formal description of nine new species. Australasian Flannery, T. F., Rich, T. H., Turnbull, W. D. and Lundelius Jr., E. Journal of Herpetology 37:48-64. L. 1992. The Macropodoidea (Marsupialia) of the early Pliocene Hoser, R. T. 2019a. 11 new species, 4 new subspecies and a Hamilton Local Fauna, Victoria. Australia. Field. Geol. 25:1-37. subgenus of Australian Dragon Lizard in the genus Gibson, L. A. and McKenzie, N. L. 2012. Occurrence of non- Tympanocryptis Peters, 1863, with a warning on the volant mammals on islands along the Kimberley coast of conservation status and long-term survival prospects of some Western Australia. pp. 15-39. in: Gibson, L. A., Yates, S. and newly named taxa. Australasian Journal of Herpetology 39:23- Doughty, P. (eds), Biodiversity values on selected Kimberley 52. islands, Australia. Western Australian Museum Records Hoser, R. T. 2019b. Richard Shine et al. (1987), Hinrich Kaiser et Supplement No. 81. Western Australian Museum, Perth, al. (2013), Jane Melville et al. (2018 and 2019): Australian Western Australia, Australia. Agamids and how rule breakers, liars, thieves, taxonomic Goodfellow, D. 1993. Fauna of Kakadu and the Top End. vandals and law breaking copyright infringers are causing reptile Wakefield Press. species to become extinct. Australasian Journal of Herpetology Gould, J. 1841. On two new species of kangaroos (Macropus 39:53-63. manicatus and M. brachyotis) from Swan River. Proceedings of Iredale, T. and Troughton, E. L. G. 1934. A check-list of the the Zoological Society of London 1840:127-128. mammals recorded from Australia. Australian Museum Memoir Gould, J. 1842. On two new species of kangaroo (Petrogale 6:1-122. [4 May 1934]. concinna and Halmaturus Binoe). Proceedings of the Zoological Kitchener, D. J. and Sanson, G. 1978. Petrogale burbidgei Society of London 1842:57. (Marsupialia, Macropodidae), a new rock wallaby from the Gray, J. E. 1837. Description of some new or little known Kimberley, Western Australia. Records of the Western Mammalia, principally in the British Museum Collection. The Australian Museum 6:269-285. Magazine of Natural History, n. s. 1:577-587. Lawlor, T. E. 1979. Handbook to the Orders and Families of Groves, C. P., Wilson, D. E. and Reeder, D. M. (eds.) 2005. Living Mammals. Mad River Press, Inc. Eureka, California, USA. Mammal Species of the World: A Taxonomic and Geographic Le Souef, A. S. 1924. Notes on some rock wallabies, genus Reference (3rd ed.). Johns Hopkins University Press, Baltimore, Petrogale, with descriptions of two new species. Australian USA. Zoologist 3:272-276. Hoser, R. T. 1991. Endangered Animals of Australia. Pierson Maynes, G. M. 1982. A new species of rock wallaby, Petrogale Publishing, Mosman, NSW, Australia:240 pp. persephone (Marsupialia: Macropodidae), from Proserpine, Hoser, R. T. 1993. Smuggled: The Underground Trade in central Queensland. Australian Mammalogy 5:47-58. Australia’s Wildlife. Apollo, Mosman, NSW, Australia. 160 pp. Maxwell, S., Burbidge, A. A. and Morris, K. 1996. The 1996 Hoser, R. T. 1996. Smuggled-2: Wildlife trafficking, crime and Action Plan for Australian Marsupials and Monotremes. corruption in Australia. Kotabi Publishing, Doncaster, Victoria, Australasian Marsupial and Monotreme Specialist Group, IUCN Australia:280 pp. Species Survival Commission, Gland, Switzerland. Hoser, R. T. 2010. Sam the scam: Sam the Koala is an Menkhorst, P. 2001. A Field Guide to the Mammals of Australia. imposter. Australasian Journal of Herpetology 8:1-64. Oxford University Press. Hoser, R. T. 2014. The break up of Odatria (Kimberleyvaranus) Meredith, R. W., Westerman, M. and Springer, M. S. 2008. A glebopalma (Mitchell, 1955) into three obvious subspecies. phylogeny and timescale for the living genera of kangaroos and Australasian Journal of Herpetology 24:16-19. kin (Macropodiformes: Marsupialia) based on nuclear DNA sequences. Aust. J. Zool. 56:395-410. Hoser, R. T. 2015. Australian agamids: Eighteen new species from the genera Amphibolurus Wagler, 1830, Lophognathus Pearson, D. 2012. Recovery plan for five species of rock Gray, 1842, Rankinia Wells and Wellington, 1984, Diporiphora wallabies: Black-footed rock wallaby (Petrogale lateralis) Short- Gray, 1842, Tympanocryptis Peters, 1863, as well as three new eared rock wallaby (Petrogale brachyotis), Monjon (Petrogale genera and six new subgenera. Australasian Journal of burbidgei), Nabarlek (Petrogale concinna), Rothschild rock Herpetology 30:37-64. wallaby (Petrogale rothschildi). Wildlife Management Program

No. 55. Department of Environment and Conservation, Perth, Felis catus (Carnivora: Felidae) on an isolated rock-wallaby Western Australia, Australia. colony in tropical Queensland. Australian Mammalogy 14:143- Potter, S., Eldridge, M. D. B., Taggart, D. A., and Cooper, S. J. 144. B. 2012a. Multiple biogeographic barriers identified across the Start, A. N., Burbidge, A. A., McDowell, M. C. and McKenzie, N. monsoon tropics of northern Australia: phylogeographic analysis L. 2012. The status of non-volant mammals along a rainfall of the brachyotis group of rock-wallabies. Molecular Ecology gradient in the south-west Kimberley, Western Australia. 21:2254-2269. Australian Mammalogy 34:36-48. Potter, S., Cooper, S. J. B., Metcalfe, C. J., Taggart, D. A., and Strahan, R. (ed.) 1988. The Australian Museum Complete book Eldridge, M. D. B. 2012b. Phylogenetic relationships within of Australian Mammals / The National Photographic Index of Petrogale (Marsupialia: Macropodidae) and their biogeographic Australian Wildlife. Angus and Robertson, North Ryde, NSW, history within Australia. Molecular Phylogenetics and Evolution Australia:530 pp. 62: 640-652. Telfer, W. R. and Eldridge, M. D. B. 2010. High levels of Potter, S., Eldridge, M. D. B., Cooper, S. J. B., Paplinska, J. Z. mitochondrial DNA divergence within short-eared rock-wallaby and Taggart, D. A. 2012c. Habitat connectivity, more than (Petrogale brachyotis) populations in northern Australia. species’ biology, influences genetic differentiation in a habitat Australian Journal of Zoology 58:104-112. specialist, the short-eared rock-wallaby (Petrogale brachyotis). Thomas, O. 1888. Catalogue of the Marsupialia and Conservation Genetics 13:937-952. Monotremata in the Collection of the British Museum (Natural Potter, S., Close, R. L., Taggart, D. A., Cooper, S. J. B. and History). British Museum, London, UK:xiii+401 pp+28 pl. Eldridge, M. D. B. 2014. Taxonomy of rock-wallabies, Petrogale Thomas, O. 1904a. On a collection of mammals made by Mr J. (Marsupialia: Macropodidae). IV. Multifaceted study of the T. Tunney in Arnhemland, Northern Territory of South Australia. brachyotis group identifies additional taxa. Australian Journal of Novitates Zoologicae 11:222-229. Zoology, 2014, 62, 401-414. Thomas, O. 1904b. On a new rock-wallaby from north-west Ride, W. D. L. 1970. A Guide to the Native Mammals of Australia. Novitates Zoologicae 11:365-366. Australia. Oxford University Press. Thomas, O. 1909. Two new mammals from N. Australia. Ann. Ride, W. D. L. (ed.) et al. (on behalf of the International Mag. Nat. Hist. 8(4):197-198. Commission on Zoological Nomenclature) 1999. International Thomas, O. 1922. A selection of lectotypes of the typical code of Zoological Nomenclature. The Natural History Museum - Australian marsupials in the British Museum Collection. Ann. Cromwell Road, London SW7 5BD, UK (also commonly cited as Mag. Nat. Hist. 9(10):127-128. “The Rules”, “Zoological Rules” or “ICZN 1999”). Thomas, O. 1926a. Two new rock-wallabies (Petrogale) Roache, M. 2011. The action plan for threatened Australian discovered by Capt. G. W. Wilkins in Northern Australia. Annals macropods. WWF-Australia, Sydney, Australia. & Magazine of Natural History 9(17):184-186. Saunders, S. 2019. Morrison and Shorten’s Big Australia: The Thomas, O. 1926b. On rock-wallabies of the Petrogale overpopulation problem. News article published 17 April 2019 penicillata-assimilis group. Annals & Magazine of Natural History and downloaded from: (9)17: 628-629. https://independentaustralia.net/politics/politics-display/morrison- Vaughan, T. A. 1986. Mammalogy, 3rd Edition. Harcourt Brace and-shortens-big-australia-the-overpopulation-problem,12584 Jovanovich College Publishers. New York, USA. Sharman, G. B., Close, R. L. and Maynes, G. M. 1990. Zaczek, Z. 2019. Former PM Kevin Rudd says Australia should Chromosome evolution, phylogeny and speciation of rock drastically increase its population to 50 MILLION to make sure wallabies (Petrogale: Macropodidae). Australian Journal of nation can defend itself in the face of threats from China. Daily Zoology 37:351-363. Mail (Australia) (27 November), published online at: https:// Shuker, K. and Durrell, G. 1993. Lost Ark: New and www.dailymail.co.uk/news/article-7729689/Kevin-Rudd-says- Rediscovered Animals of the Twentieth Century. Harper Collins Australia-increase-population-50-MILLION-defend-China.html Publishers, New York, USA. CONFLICTS OF INTEREST Spencer, P. B. S. 1991. Evidence of predation by a feral cat, None.

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Hoser 2020 - Australasian Journal of Herpetology 42:42-49. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 50 Australasian Journal of Herpetology Australasian Journal of Herpetology 42:50-64. Published 25 April 2020. ISSN 1836-5698 (Print) ISSN 1836-5779 (Online)

An overdue refinement of the taxonomy of the Australian Ring Tailed Dragons, Genus Ctenophorus Fitzinger, 1843, Subgenus Tachyon Wells and Wellington, 1985, including the formal descriptions of eight new species.

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RAYMOND T. HOSER

LSID urn:lsid:zoobank.org:author:F9D74EB5-CFB5-49A0-8C7C-9F993B8504AE

488 Park Road, Park Orchards, Victoria, 3134, Australia. Phone: +61 3 9812 3322 Fax: 9812 3355 E-mail: snakeman (at) snakeman.com.au Received 10 December 2019, Accepted 12 March 2020, Published 25 April 2020.

ABSTRACT The Australian Ring-tailed Dragons of the Genus Ctenophorus Fitzinger, 1843, Subgenus Tachyon Wells and Wellington, 1985 as defined by Hoser in 2015, have been generally neglected in their taxonomy since the publication of Wells and Wellington (1985). Those authors formally divided the species Grammatophora caudicincta Günther, 1875 six ways in line with the earlier subspecies divisions of Storr (1967) and placed all into their newly created genus Tachyon. This arrangement was vehemently opposed by a group of pseudo-scientists known as the Wolfgang Wüster gang for reasons of personal animosity rather than science. Furthermore due to the ruthless and improper methods of the group (as demonstrated in a war-cry document called Kaiser et al. 2013), they have effectively forced other publishing herpetologists to refuse to accept the Wells and Wellington taxonomy and nomenclature in the three and half decades since the 1985 paper of Wells and Wellington. In spite of lies, deception and so-called smoke and mirrors as practiced by the Wolfgang Wüster gang, science progresses and in line with this, Melville et al. (2016) not only broadly validated the much lampooned taxonomy and nomenclature of Wells and Wellington (1985), but furthermore wholly validated the taxonomy of Hoser in 2015. Melville et al. (2016) also provided sound evidence of the presence of at least seven more unnamed species in the complex, all diverged from nearest ancestors in the Pliocene Epoch (at least 2.5 MYA) mirroring the morphological evidence of Storr (1967). This paper in effect combines the results of Storr (1967) with Melville et al. (2016) with the added benefit of inspection of live specimens from the seven relevant populations and all previously named forms to formally describe and name eight new species in accordance with the rules of the International Code of Zoological Nomenclature (Ride et al. 1999). The eighth named species is separated from others by a biogeographical barrier of known antiquity. Each is estimated to have diverged from their nearest common ancestor at least 2.5 million years prior and as they are ecologically and distributionally separated, are clearly separate species by any currently used definition. The first ever proper diagnosis of the expanded subgenus Tachyon Wells and Wellington, 1985 was published by Hoser in 2015. It has stood the test of time and is repeated in this paper. Keywords: Taxonomy; nomenclature; lizards; dragons; Agamidae; Ctenophorus; Tachyon; Grammatophora; Amphibolurus; Western Australia; Queensland; Northern Territory; Australia; Ring-tailed dragon; caudicinctus; caudicincta; yinnietharra; graafi; imbricatus; infans; macropus; mensarum; slateri; new species; adelynhoserae; jackyhoserae; katrinahoserae; lenhoseri; maxinehoserae; ronhoseri; sharonhoserae; shireenhoserae.

INTRODUCTION unnamed) were examined both live in the wild and via museum As part of an ongoing audit of Australia’s reptiles and frogs, the collections and/or their records, including all State and Territory lizards within the putative genus Tachyon Wells and Wellington, Museums on mainland Australia. Furthermore photos and data 1985, (herein treated as a subgenus), better known as the with accurate locality data was also assessed, as was all relevant Australian Ring Tailed Dragons were examined with a view to previously published scientific literature and the so-called grey confirming the taxonomy and nomenclature of relevant species or literature in the form of popular mass-market books, internet subspecies as being correct, or in the alternative being altered to sites, blogs, photo-sharing sites and the like. reflect the biological reality. The final results of this audit found that within the so-called Tachyon was originally erected as a genus by Wells and Grammatophora caudicincta Günther, 1875 complex, there were Wellington (1985), but the molecular evidence of Pyron et al. eight recognized species, these being those six cited by Wells (2013) suggested that a more accurate placement of the relevant and Wellington (1985) on page 20 at top, as well as the species species was as a subgenus within the better-known Ctenophorus originally described as Grammatophora ornata Gray, 1854, and Fitzinger, 1843. also the species originally described as Amphibolurus Hoser (2015g) was the first publishing herpetologist since Wells yinneatharra Storr, 1981, both of which Wells and Wellington and Wellington (1985) to utilize the genus name Tachyon, but in (1985) placed in the genus Ctenophorus Fitzinger, 1843 (type line with the results of Pyron et al. (2013) relegated the genus to species Grammatophora decresii Duméril and Bibron, 1837). a subgenus, within the genus Ctenophorus. The genus was also There were also eight more forms within the subgenus Tachyon expanded to include two closely related species, namely C. worthy of species-level recognition based on molecular yinnietharra (Storr, 1981) and C. ornatus (Gray, 1845). divergence, morphological differences and geographical Hoser (2015g) treated all other previously named forms of the disjunction caused by well-established biogeographical barriers three putative species as subspecies, all within C. caudicincta of significant and known antiquity. (Günther, 1875) pending the publication of this paper. In summary the relevant unnamed species are as follows: Specimens of all relevant species or subspecies (named and until 1/ The species Ctenophorus macropus (Storr, 1967) of northern now unnamed) were examined both live in the wild and via Australia was found to comprise five geographically separated museum collections and their records, including all State and species. Each was restricted to relevant rock formations Territory Museums on mainland Australia. Furthermore photos (mountain ranges and outliers) and separated from one another and data with accurate locality data was also assessed, as was by relatively flat intervening areas, which happen to be well- all relevant previously published scientific literature and the so- known biogeographical barriers of known antiquity. called grey literature in the form of popular mass-market books, Type C. macropus is the form from Arnhem Land in the Northern internet sites, blogs, photo-sharing sites and the like. Territory, while two populations in the Kimberley Ranges of Two papers of key relevance to the relevant taxa were those of Western Australia, one from the Gulf of Carpentaria and one from Storr (1967), which had a detailed morphological analysis of most the Selwyn Ranges, north-west Queensland (around Mount Isa relevant species and/or subspecies, as well as a more recent and south of there) were unnamed. These are formally described paper of Melville et al. (2016), which inspected the same putative herein. taxa at the molecular level, including the species originally 2/ The species C. slateri (Storr, 1967) is clearly composite, with described as Grammatophora ornata Gray, 1854, which clearly the nominate form from Central Australia and the northern fell within the broader species grouping and the putative genus population is unnamed and so it is formally described herein as a Tachyon Wells and Wellington, 1985 as defined by Hoser new species. (2015g). 3/ Nominate C. caudicincta (Günther, 1875) including as I note that Melville et al. (2016) failed to inspect the relevant identified by Melville et al. (2016) with a distribution centred on species C. yinneatharra Storr, 1981, although it’s position in their the Pilbara region of Western Australia was shown by them to published phylogeny could be easily inferred as being closest to consist of two allopatric species and the previously unnamed their clade labelled as “C. ornatus”. They also failed to inspect south-east Pilbara form is formally described herein. specimens from the Mount Isa area or south of there, even 4/ The morphologically distinct and genetically divergent though they form a separate biogeographically isolated population of putative Grammatophora ornata Gray, 1854 (herein population, apart from those they inspected. placed within the subgenus Tachyon) from the north-west part of The combined evidence of these papers alone gave rise to a the range of the putative species distribution is herein described well-founded belief that there were at least seven unnamed forms as a new species as is another divergent form from the inland at the species level. parts of the south-east of Western Australia (the Goldfields Relevant specimens were examined and confirmed that each of region). these forms warranted recognition at the species level, which is Gray’s holotype specimen, lacks specific locality data. the main basis for publishing this paper. That is to formally name But based on the original written descriptions of Gray and and make available names for the seven species-level taxa, and Boulenger (1885) it is clearly of the form from near Perth, the other eighth species-level taxon not inspected by either Western Australia and so it is the nominate species C. ornata. Melville et al. (2016) or Storr (1967), all being named in In passing I note that the statement by Melville et al. (2016), accordance with the rules of the International Code of Zoological “With our recommendations the C. caudicinctus species group, Nomenclature (Ride et al. 1999). which currently incorporates six subspecies, would become four MATERIALS, METHODS AND RESULTS species: C. caudicinctus, C. infans, C. slateri and C. graafi.” did These are inferred in both the abstract and introduction and self not make any sense at all, even when reconciled exclusively with evident in the descriptions that follow. the data the same authors presented in the very same paper and noting that the authors also recklessly overlooked the species C. An audit of relevant species of Australian Ring-tailed Dragons of yinneatharra (Storr, 1981), even though a year prior Hoser the subgenus Tachyon Wells and Wellington, 1985 sensu lato as (2015g) had correctly placed that taxon within the species group. defined by Hoser (2015g) confirmed the generic level assignment It is self-evident from the morphological evidence of Storr (1967) of species and validity of the relevant named forms as identified and molecular evidence of Melville et al. (2016) that the case for by Wells and Wellington (1985) as placed by Hoser (2015g) and/ recognition of C. mensarum (Storr, 1967) as a full species is or in line with it, as well as the species originally described as weaker than for all other species (including the five formally Grammatophora ornata Gray, 1854, which is clearly a part of the named herein). species complex (= same subgenus) and C. yinneatharra (Storr, 1981), as previously allocated by Hoser (2015g). This includes significantly less molecular divergence from nominate C. caudicincta (as shown by Melville et al. 2016) than Specimens of all relevant species (named and until now Hoser 2020 - Australasian Journal of Herpetology 42:50-64. for all other putative species, including the eight formally named Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 52 Australasian Journal of Herpetology herein. 2020, unless otherwise stated and was accurate in terms of the However that taxon C. mensarum (Storr, 1967) is tentatively context cited herein as of that date. recognized at the full species level within this paper pending Unless otherwise stated explicitly, colour descriptions apply to further work on allegedly intermediate specimens between the living adult male specimens of generally good health and not two putative forms. under any form of stress by means such as excessive cool, heat, The genus-level arrangement of relevant species and other dehydration or abnormal skin reaction to chemical or other input. Australian agamids was found to be in accordance with the While numerous texts and references were consulted prior to published results of Hoser (2015g), which was so accurate as to publication of this paper, the criteria used to separate the relevant not needing any alteration of diagnoses at this point in time. species has already been spelt out and/or is done so within each Newly named species herein simply fit within the same diagnosed formal description and does not rely on material within genus or subgenus as does their nearest known relative. publications not explicitly cited herein. The literature relevant to the taxonomy and nomenclature of the Each newly named species is readily and consistently separable subgenus Tachyon as first defined by Wells and Wellington from their nearest congener and that which until now it has been (1985) and redefined by Hoser (2015g) and herein, including the previously treated as. taxonomic and nomenclatural decisions herein include the Delays in recognition of these species could jeopardise the long- following: Ahl (1926), Baverstock and Bradshaw (1975), term survival of these taxa as outlined by Hoser (2019a, 2019b) Boulenger (1885), Bradshaw (1970, 1971), Bradshaw and Main and sources cited therein. (1968), Bradshaw and Shoemaker (1967), Cogger (2014), Therefore attempts by taxonomic vandals like the Wolfgang Cogger et al. (1983), Denzer et al. (1997), Doody and Schembri Wüster gang via Kaiser (2012a, 2012b, 2013, 2014a, 2014b) and (2014), Even (2005), Fitzinger (1843), Glauert (1959), Gray Kaiser et al. (2013) (as frequently amended) to unlawfully (1845), Günther (1875), Hoser (2015g), Kinghorn (1924), Lebas suppress the recognition of these taxa on the basis they have a and Spencer (2000), Melville et al. (2001, 2016), Peters (1876), personal dislike for the person who formally named it should be Pyron et al. (2013), Ride et al. (1999), Sternfeld (1925), Storr resisted (Dubois et al. 2019). (1967, 1981), Storr et al. (1983), Wells and Wellington (1985) Claims by the Wüster gang against this paper and the Wilson and Knowles (1988), Wilson and Swan (2017) and descriptions herein will no doubt be no different to those the gang sources cited therein. have made previously, all of which were discredited long ago as FURTHER DISCUSSION RELEVANT TO THIS PUBLICATION outlined by Dubois et al. (2019), Hoser, (2007, 2009, 2012a, An illegal armed raid and theft of materials on 17 Aug 2011 2012b, 2013a, 2015a-f, 2019a, 2019b) and sources cited therein. effectively stopped the publication of a variant of this paper being The following genus and subgenus-level diagnosis of the relevant published back then and a significant amount of materials taken lizard species is taken in an abridged form from Hoser (2015g) in that raid was not returned. This was in spite of court orders and excluding formal diagnoses of the subgenera not subject of telling the relevant State Wildlife officers to do so (Court of this paper. Appeal 2014, Victorian Civil and Administrative Tribunal 2015). Formal descriptions of the eight relevant species follow. Rather than run the risk of species becoming threatened or Information relevant to conservation of Australian reptiles in extinct due to non-recognition of them as shown in Hoser (2019a, Hoser (1989, 1991, 1993 and 1996) and relevant comments in 2019b), I have instead opted to publish this paper in its current Hoser (2019a, 2019b) applies to the newly named taxa herein. form, even though a significant amount of further data was GENUS CTENOPHORUS FITZINGER, 1843. intended to be published and is not. Type species: Grammatophora decresii Dumeìril and Bibron Naming of taxa is perhaps the most important step in their 1837. ultimate preservation and it is with this motivation in mind Diagnosis: Ctenophorus as defined until now (see also Cogger (protection of biodiversity) that I have chosen to publish this 2014) is defined by the following definition, modified to take into paper. account the genera defined by Hoser (2015g) being the most Until now, no new (and generally recognized) taxa within the so- recent full and proper treatment of the genus. Ctenophorus is called Grammatophora caudicincta Günther, 1875 complex of defined as an Australian agamid genus characterised by small species has been formally identified or named since the paper of dorsal scales, homogenous or with at most slightly enlarged Storr (1967). tubercles; a few species with distinct rows of paravertebral or INFORMATION RELEVANT TO THE FORMAL DESCRIPTIONS dorsolateral spinose scales; a row of enlarged scales from below THAT FOLLOW the eye to above the ear; tympanum exposed (not exposed in There is no conflict of interest in terms of this paper or the Notactenophorus Hoser 2015) and most Pseudoctenophorus conclusions arrived at herein. Hoser, 2015; tail long, ranging from slightly to much longer than Several people including anonymous peer reviewers who revised the head and body; femoral and preanal pores present in males; the manuscript prior to publication are also thanked as are adult males usually with distinctive black or dark grey markings relevant staff at museums who made specimens and records on the throat and/or chest. available in line with international obligations. The genus Paractenophorus Hoser, 2015 is separated from In terms of the following formal descriptions, spellings should not Ctenophorus, Notactenophorus Hoser 2015 and be altered in any way for any purpose unless expressly and Pseudoctenophorus Hoser, 2015 by the following suite of exclusively called for by the rules governing Zoological characters: tympanum exposed; no series of enlarged, spinose Nomenclature as administered by the International Commission scales on either side of the base of the tail; hind limb reaching no of Zoological Nomenclature. further than the tympanum when adpressed; tail usually less than In the unlikely event two newly named taxa are deemed 1.5 times as long as the head and body; nasal region is not conspecific by a first reviser, then the name to be used and swollen, the nostril lying below an angular canthal ridge; pores retained is that which first appears in this paper by way of page fewer than 15; nostril is slit-like or narrowly elliptical. priority and as listed in the abstract keywords. Specimens within the genus Pseudoctenophorus Hoser, 2015 Some material in descriptions for taxa may be repeated for other are separated from all other Ctenophorus Fitzinger, 1843, the taxa in this paper and this is necessary to ensure each fully genus they were placed in previously, by the following suite of complies with the provisions of the International Code of characters, being one or other of the following three: Zoological Nomenclature (Fourth edition) (Ride et al. 1999) as 1/ Tympanum exposed; a series of enlarged, spinose scales on amended online since. either side of the base of the tail (subgenus Pseudoctenophorus Material downloaded from the internet and cited anywhere in this Hoser, 2015), or: paper was downloaded and checked most recently as of 1 March 2/ Tympanum hidden; covered by skin; body scales are strongly Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 53

heterogeneous, many of the larger scales on the body and head a later scientist is forced to waste their time and correct the keeled or moderately spinose; a series of enlarged, spinose historical record and re-instate the correct Wells and Wellington scales on either side of the base of the tail (subgenus nomen. Chapmanagama Hoser, 2015), or: The time spent doing this would be better diverted towards new 3/ Tympanum hidden; covered by skin; body scales are strongly science and the wildlife conservation objectives that this serves. heterogeneous, many of the larger scales on the body and head This is particularly the case in the context of Australian reptiles, keeled or moderately spinose; no series of enlarged, spinose where as of 2020 dozens of species still await formal description. scales on either side of the base of the tail (subgenus SUBGENUS TACHYON WELLS AND WELLINGTON, 1985. Turnbullagama Hoser, 2015). Type species: Grammatophora caudicincta Günther, 1875. Notactenophorus Hoser, 2015 is readily separated from all other Diagnosis: Species within the subgenus Tachyon Wells and members of the genus Ctenophorus (where it has been placed Wellington, 1985 are separated from all other Ctenophorus until now, as defined in Cogger 2014) and Pseudoctenophorus Fitzinger, 1843 by the following suite of characters being one or Hoser, 2015, by the following unique suite of characters: other of: Tympanum is hidden being covered by skin, the body scales are 1/ Tympanum exposed; no series of enlarged, spinose scales on smooth, mostly small, homogenous, with scattered larger but either side of the base of the tail; hind limb usually reaching to small, flat scales, not keeled or spinose, with a dorsal pattern of a eye or beyond when adpressed; tail usually much more than 1.5 longitudinal dorso-lateral series of five or six large black spots on times as long as the head and body; canthus rostralis swollen, either side. but nostrils, when viewed from above, face distinctly upwards as Distribution: Most parts of continental Australia. opposed to outwards (species C. adelynhoserae sp. nov.; C. Content: C. decresii (Duméril and Bibron, 1837) (type species); caudicinctus (Günther, 1875); C. graafi (Storr, 1967); C. C. adelynhoserae sp. nov.; C. caudicinctus (Günther, 1875); C. imbricatus (Peters, 1876); C. infans (Storr, 1967); C. cristatus (Gray, 1841); C. dudleyi Wells and Wellington 1985; C. jackyhoserae sp. nov.; C. katrinahoserae sp. nov.; C. lenhoseri femoralis (Storr, 1965); C. fionni (Procter, 1923); C. fordi (Storr, sp. nov.; C. macropus (Storr, 1967); C. maxinehoserae sp. nov.; 1965); C. gibba (Houston, 1974); C. graafi (Storr, 1967); C. C. mensarum (Storr, 1967); C. ronhoseri sp. nov.; C. slateri hawkeswoodi (Wells and Wellington, 1985); C. imbricatus (Storr, 1967)) or: (Peters, 1876); C. infans (Storr, 1967); C. isolepis (Fischer, 2/ Tympanum exposed; no series of enlarged, spinose scales on 1881); C. jackyhoserae sp. nov.; C. katrinahoserae sp. nov.; C. either side of the base of the tail; hind limb usually reaching to lenhoseri sp. nov.; C. macropus (Storr, 1967); C. maculatus eye or beyond when adpressed; tail usually much more than 1.5 (Gray, 1831); C. maxinehoserae sp. nov.; C. mckenziei (Storr, times as long as the head and body; canthus rostralis angular or 1981); C. mirrityana McLean, Moussalli, Sass and Stuart-Fox, moderately swollen, but nostrils, when viewed from above, face 2013; C. nguyarna Doughty, Maryan, Melville and Austin, 2007; outwards as opposed to distinctly upwards (as seen in the C. mensarum (Storr, 1967); C. nuchalis (De Vis, 1884); C. species C. adelynhoserae sp. nov.; C. caudicinctus (Günther, ornatus (Gray, 1845); C. pictus (Peters, 1866); C. raffertyi Wells 1875); C. graafi (Storr, 1967); C. imbricatus (Peters, 1876); C. and Wellington, 1985; C. reticulatus (Gray, 1845); C. ronhoseri infans (Storr, 1967); C. jackyhoserae sp. nov.; C. katrinahoserae sp. nov.; C. rubens (Storr, 1965); C. rufescens (Stirling and Zietz, sp. nov.; C. lenhoseri sp. nov.; C. macropus (Storr, 1967); C. 1893); C. salinarum Storr, 1966; C. scutulatus (Stirling and Zietz, maxinehoserae sp. nov.; C. mensarum (Storr, 1967); C. 1893); C. sharonhoserae sp. nov.; C. shireenhoserae sp. nov.; ronhoseri sp. nov.; C. slateri (Storr, 1967)); at most a few C. slateri (Storr, 1967); C. tjantjalka Johnston, 1992; C. enlarged keeled scales on the nape; a series of enlarged vadnappa Houston, 1974; C. yinnietharra (Storr, 1981). vertebral scales, if present, forming a distinct linear series only to Comments: All of C. graafi (Storr, 1967); C. imbricatus (Peters, about the level of the forelimbs; dorsal scales at most with low, 1876); C. infans (Storr, 1967); C. macropus (Storr, 1967); C. irregular keels which do not form distinct continuous ridges; mensarum (Storr, 1967); C. slateri (Storr, 1967) were treated by dorsolateral scales and those on the chest smooth, or with low Hoser (2015g) as synonyms of C. caudicinctus (Günther, 1875) blunt edges; nostril elliptical in a swollen nasal scale lying on a pending publication of this paper. swollen canthal ridge; tibial region with a series of anterior Hoser (2015g) erroneously treated C. raffertyi Wells and proximal scales which are very much larger than those on the Wellington, 1985 as a synonym of C. clayi (Storr, 1967) by posterior surface (species C. shireenhoserae sp. nov.; C. overlooking it in an error also not picked up in peer review. sharonhoserae sp. nov.; C. ornatus and C. yinnietharra). However the taxon C. raffertyi is herein regarded as valid on the Ctenophorus as defined until now (Cogger 2014, Hoser 2015g) is basis of morphological and distributional divergence across a defined by the following definition, modified to take into account biogeographical barrier of known antiquity. the new genera as defined herein. Ctenophorus is defined as an In an act of taxonomic vandalism, Sadlier et al. (2019) Australian agamid genus characterised by small dorsal scales, deliberately illegally renamed the taxon C. hawkeswoodi (Wells homogenous or with at most slightly enlarged tubercles; a few and Wellington, 1985) as recognized as valid by Hoser (2015g) species with distinct rows of paravertebral or dorsolateral spinose as Ctenophorus spinodomus Sadlier et al. (2019). scales; a row of enlarged scales from below the eye to above the The International Code of Zoological Nomenclature (Ride et al. ear; tympanum exposed (not exposed in Notactenophorus Hoser, 1999) is clear in its directives in terms of the rule of priority and so 2015 and most Pseudoctenophorus Hoser, 2015); tail long, the correct name for this taxon is C. hawkeswoodi (Wells and ranging from slightly to much longer than the head and body; Wellington, 1985). femoral and preanal pores present in males; adult males usually That species is part of the C. fordi (Storr, 1965) species complex with distinctive black or dark grey markings on the throat and/or subject of another paper (Hoser 2000). chest. While it is self-evident that some of the species diagnoses in the The genus Paractenophorus Hoser, 2015 is separated from paper of Wells and Wellington (1985) are vague and imprecise, Ctenophorus, Notactenophorus Hoser, 2015 and the fact remains that the relevant names are available in terms of Pseudoctenophorus Hoser, 2015 by the following suite of the rules of the International Code of Zoological Nomenclature, characters: tympanum exposed; no series of enlarged, spinose meaning that the relevant paper should be consulted if and when scales on either side of the base of the tail; hind limb reaching no herpetologists are doing taxonomic works on Australian reptiles further than the tympanum when adpressed; tail usually less than and frogs and there is a likelihood of a priority name being 1.5 times as long as the head and body; nasal region is not available for a taxon. swollen, the nostril lying below an angular canthal ridge; pores Overwriting a Wells and Wellington name may give a later worker fewer than 15; nostril is slit-like or narrowly elliptical. a sense of greatness in being able to claim “discovery” of a Specimens within the genus Pseudoctenophorus Hoser, 2015

Hoser 2020 - Australasian Journal of Herpetology 42:50-64. species, but this “discovery” will evaporate at a future date when are separated from all other Ctenophorus Fitzinger, 1843, the Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 54 Australasian Journal of Herpetology genus they were placed previously, by the following suite of not inspected by Melville et al. (2016). characters, being one or other of the following three: C. lenhoseri sp. nov. is however separated from the other species 1/ Tympanum exposed; a series of enlarged, spinose scales on by a barrier of known antiquity in far north-west Queensland. either side of the base of the tail (subgenus Pseudoctenophorus Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. nov., Hoser, 2015), or: C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. 2/ Tympanum hidden; covered by skin; body scales are strongly macropus are readily separated by their unique colour patterns. heterogeneous, many of the larger scales on the body and head Adult male Ctenophorus adelynhoserae sp. nov. are separated keeled or moderately spinose; a series of enlarged, spinose from males of the other species by having a brown dorsum and scales on either side of the base of the tail (subgenus flanks with either no markings or indistinct ones. Legs are either Chapmanagama Hoser, 2015), or: unmarked or lack obvious markings, which otherwise may include 3/ Tympanum hidden; covered by skin; body scales are strongly indistinct spots or crossbands. There are scattered dark flecks or heterogeneous, many of the larger scales on the body and head peppering along the lower sides and no light blue spots or flecks keeled or moderately spinose; no series of enlarged, spinose anywhere on the dorsum. Between the eye and ear is a dark and scales on either side of the base of the tail (subgenus semi-distinct bar or broken bar. There are 19-21 tail rings, being Turnbullagama Hoser, 2015). alternating brown and whitish, the lighter sections being narrower Notactenophorus Hoser, 2015 is readily separated from all other than the darker sections. members of the genus Ctenophorus (where it has been placed Adult females are greyish in colouration and with a greyish head until now, as defined in Cogger 2014) and Pseudoctenophorus with mottling as opposed to any well-defined markings. Hoser, 2015 by the following unique suite of characters: Adult male C. jackyhoserae sp. nov. normally have a brownish- Tympanum is hidden being covered by skin, the body scales are red dorsal surface with broken lines or evenly arranged flecks smooth, mostly small, homogenous, with scattered larger but along, but not on the mid-dorsal line. Some specimens small, flat scales, not keeled or spinose, with a dorsal pattern of a alternatively have a whitish-red upper body to deep red, longitudinal dorso-lateral series of five or six large black spots on characterised by a dorsal arrangement of about six broken either side. orange-brown-red lines running longitudinally down the body from Distribution: Drier parts of northern, central and Western neck to rump, being broken by intervening areas of creamy-grey Australia, including the south-west and invariably associated with white that are wider than the darker markings. The side of the rock outcrops. head and labials are whitish or at least with whiter pigment than Content: Ctenophorus (Tachyon) caudicinctus (Günther, 1875) elsewhere and with indistinct reddish markings near the ear and (type species); C. (Tachyon) adelynhoserae sp. nov.; C. temples. The top of the head is a dark reddish orange, The lower (Tachyon) graafi (Storr, 1967); C. (Tachyon) imbricatus (Peters, flanks are characterised by 3-5 dark brown bars across a pale 1876); C. (Tachyon) infans (Storr, 1967); C. (Tachyon) yellowish background, being indistinctly divided from the whitish- jackyhoserae sp. nov.; C. (Tachyon) katrinahoserae sp. nov.; C. grey above, or in some specimens these may reduce to being (Tachyon) lenhoseri sp. nov.; C. (Tachyon) macropus (Storr, evenly spaced paired dark spots. There are no light blue spots or 1967); C. (Tachyon) maxinehoserae sp. nov.; C. (Tachyon) flecks anywhere on the dorsum. There is no dark bar between the mensarum (Storr, 1967); C. (Tachyon) ornatus (Gray, 1845); C. eye and ear, where the stripe would otherwise be seen in (Tachyon) ronhoseri sp. nov.; C. (Tachyon) sharonhoserae sp. Ctenophorus adelynhoserae sp. nov.. nov.; C. (Tachyon) shireenhoserae sp. nov.; C. (Tachyon) slateri There are 17-18 tail rings, alternating blackish and whitish, the (Storr, 1967); C. (Tachyon) yinnietharra (Storr, 1981). rings being of even thickness. CTENOPHORUS (TACHYON) ADELYNHOSERAE SP. NOV. Adult female C. jackyhoserae sp. nov. are of greyish colouration, featuring brown barring on the lower labial area, no markings, or LSID urn:lsid:zoobank.org:act:4A2B64E0-B3BA-430C-8BD2- very indistinct on all limbs and a dorsal surface featuring a DEFBAFFF48D8 mottled appearance of light and dark, but no obvious well defined Holotype: A preserved specimen at the Western Australian pattern. Museum, Perth, Western Australia, Australia, specimen number Adult male C. lenhoseri sp. nov. have a strongly reddish-brown R125200 collected from the Saint George Range, south-west forebody, becoming greyish towards the rear and tail. On either Kimberley Division of Western Australia, Australia, Latitude - side of the mid-dorsal line are about 5 pairs of evenly spaced 18.75 S., Longitude 125.15 E. This facility allows access to its semi-distinct small dark blackish brown spots of semi-rectangular holdings. shape, the longer sides being those running towards the flanks. Paratype: A preserved specimen at the Western Australian The flanks are more reddish than the mid dorsum, which is Museum, Perth, Western Australia, Australia, specimen number slightly greyish and the flanks also have scattered light blue spots R125199 collected from the Saint George Range, south-west or flecks. There are 24-28 tail rings, defined by having very Kimberley Division of Western Australia, Australia, Latitude - narrow light sections and wide darker sections, being alternating 18.75 S., Longitude 125.15 E. brownish-black and whitish bands. There are no obvious blotches Diagnosis: The species Ctenophorus adelynhoserae sp. nov. or spots on the lower flanks, or dark bar between the eye and ear, from the south-west Kimberley division of Western Australia although in some specimens there is dark peppering where the (being the St. George and Edgar Ranges, north-east of Broome, stripe would otherwise be seen in Ctenophorus adelynhoserae Western Australia), C. jackyhoserae sp. nov. from the northern sp. nov.. and eastern Kimberley division of Western Australia and Adult female C. lenhoseri sp. nov. are of similar colour to males, immediately adjacent parts of the Northern Territory, C. being reddish and without obvious pattern of any sort. The dorsal katrinahoserae sp. nov. from far north-west Queensland and the surface is effectively unicolour, save for a poorly defined zone of nearby parts of the Northern Territory, south of the Gulf of darkening along the mid vertebral line and 4-6 well-spaced pairs Carpentaria and C. lenhoseri sp. nov. from the Selwyn Ranges of small but obvious yellow spots on either side of the mid dorsal (generally around Mount Isa and south of there), in north-west line of the body. There are sometimes paired scattered dark spots Queensland, have until now all been treated as populations of C. around the neck, which typically fade in older specimens. There macropus (Storr, 1967) with a distribution centred on Arnhem are 18-22 tail rings, with lighter ones being narrow or incomplete Land, Northern Territory, with which they are morphologically and darker ones about 3-4 times wider, the colouration being most similar to. greyish orange (wider) and yellow (narrower) bands. Fore and However all are morphologically distinct, wholly allopatric and hind limbs both have indistinct bands. sufficiently divergent to be treated as full species as is seen Juvenile C. lenhoseri sp. nov. of both sexes are characterised by herein, with divergences estimated by Melville et al. (2016) to be a significant amount of dark brown and black pigment in the form in excess of 2.5 MYA for all but C. lenhoseri sp. nov. which was of flecks or mottling on the upper surfaces, which fades with age. Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 55

Adult male C. katrinahoserae sp. nov. have an orange-grey- eldest daughter, Adelyn Hoser, of Park Orchards, Victoria, brown forebody, with a distinct salmon colouration across whitish Australia in recognition of more than 20 years active work with parts of the upper forebody and head, becoming dull greyish wildlife conservation, research and education in Australia. towards the rear and tail. On either side of the mid-dorsal line are CTENOPHORUS (TACHYON) JACKYHOSERAE SP. NOV. about scattered semi-distinct dark flecks which may also appear LSID urn:lsid:zoobank.org:act:FA6270EE-615D-4F28-9113- on the mid flanks, but the dorsum is otherwise not prominently B5AB50B622FA marked. The flanks are slightly darker than the dorsum at the mid Holotype: A preserved specimen at the Western Australian flanks before becoming light again at the belly. There are 20-26 Museum, Perth, Western Australia, Australia, specimen number tail rings, defined by having narrow light sections and wide darker R113989 collected from 32 km west of El Questro Station sections, being alternating brownish-black and whitish bands. Kimberley Division of Western Australia, Australia, Latitude - There are no obvious blotches or spots on the lower flanks, or 16.02 S., Longitude 127.97 E. dark bar between the eye and ear, although in some specimens there is dark peppering where the stripe would otherwise be seen This facility allows access to its holdings. in Ctenophorus adelynhoserae sp. nov.. Paratype: A preserved specimen at the Northern Territory Adult female C. katrinahoserae sp. nov. are of similar colour to Museum, Darwin, Northern Territory, Australia, specimen number males, but generally greyish in colour with lighter yellow-cream R07018 collected from the Pentecost River Crossing on the Gibb interspaces. There is a mottled pattern across the dorsal surface, River Road, Kimberley Division of Western Australia, Australia, being most prominent at the anterior end and the sides of the Latitude -16.17 S., Longitude 127.98 E. head. Diagnosis: The species Ctenophorus jackyhoserae sp. nov. In both sexes, the legs are generally unmarked, but commonly from the northern and eastern Kimberley division of Western with indistinct blotching, particularly on the lower rear legs upper Australia and immediately adjacent parts of the Northern Territory, surface. C. adelynhoserae sp. nov. from the south-west Kimberley division of Western Australia (being the St. George and Edgar Ranges, Adult male C. macropus have a generally light orange dorsal north-east of Broome, Western Australia), C. katrinahoserae sp. colour, with scattered dark brown flecks on the head and neck, nov. from far north-west Queensland and the nearby parts of the but not on the body or legs. Prominent in this species are Northern Territory, south of the Gulf of Carpentaria and C. numerous bright aqua-blue spots scattered across the dorsal lenhoseri sp. nov. from the Selwyn Ranges (generally around surface and sides of the flanks. There are 19-21 tail rings, with Mount Isa and south of there), in north-west Queensland, have the darker sections being significantly wider than the lighter ones. until now all been treated as populations of C. macropus (Storr, The anterior tail is generally brownish orange with indistinct 1967) with a distribution centred on Arnhem Land, Northern bands, while the posterior half of the tail has fairly well defined Territory, with which they are morphologically most similar to. bands, the colours being brown and yellow-white. However all are morphologically distinct, wholly allopatric and Adult female C. macropus are similar in most respects to adult sufficiently divergent to be treated as full species as is seen female C. jackyhoserae sp. nov. but any yellow spotting on the herein, with divergences estimated by Melville et al. (2016) to be back near the mid-dorsal line is relatively indistinct and the front in excess of 2.5 MYA for all but C. lenhoseri sp. nov. which was legs have well defined bands. Any markings on the rear legs, if not inspected by Melville et al. (2016). present are indistinct. C. lenhoseri sp. nov. is however separated from the other Ctenophorus adelynhoserae sp. nov. in life is depicted in Storr et species by a barrier of known antiquity in far north-west al. (1983), plate 3, image 4, adult male. Queensland. C. jackyhoserae sp. nov. is depicted in life in Wilson and Swan Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. nov., (2017) on page 413 at top left, adult male and Wilson and C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. Knowles (1988), page 207, top left adult male, top right adult macropus are readily separated by their unique colour patterns. female and in Hoser (1989) page 67 top, male. Adult male Ctenophorus adelynhoserae sp. nov. are separated C. katrinahoserae sp. nov. is depicted in life online at: from males of the other species by having a brown dorsum and https://pbase.com/gehyra/image/129646530 flanks with either no markings or indistinct ones. Legs are either (last downloaded on 1 March 2020). unmarked or lack obvious markings, which otherwise may include C. lenhoseri sp. nov. is depicted in life in Cogger (2014), bottom indistinct spots or crossbands. There are scattered dark flecks or left, adult male (aged) and Brown (2014), page 653 (3 images peppering along the lower sides and no light blue spots or flecks labelled as Ctenophorus caudicinctus macropus, Windorah, anywhere on the dorsum. Between the eye and ear is a dark and Queensland, two adult males and an adult female). semi-distinct bar or broken bar. There are 19-21 tail rings, being All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. alternating brown and whitish, the lighter sections being narrower nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. than the darker sections. macropus can be separated from all congeners in the subgenus Adult female C. adelynhoserae sp. nov. are greyish in colouration Tachyon Wells and Wellington, 1985 (as defined elsewhere in and with a greyish head with mottling as opposed to any well- this paper), by having over 32 lamellae under the fourth toe defined markings. versus less than 32 in all other species as well as a tail 2 times Adult male C. jackyhoserae sp. nov. normally have a brownish- the length of snout-vent, versus less than 1.8 times snout-vent in red dorsal surface with broken lines or evenly arranged flecks all other species and the adpressed hind-leg extends past the along, but not on the mid-dorsal line. Some specimens snout (not so in all other species). alternatively have a whitish-red upper body to deep red, All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. characterised by a dorsal arrangement of about six broken nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. orange-brown-red lines running longitudinally down the body from macropus are characterised as follows: Distal three-quarters of neck to rump, being broken by intervening areas of creamy-grey tail compressed. Nasal small, located on top of obtuse rostral white that are wider than the darker markings. The side of the ridge. Keels of dorsal scales moderately strong and sharp, head and labials are whitish or at least with whiter pigment than terminating in a blunt end or short spine. Ventrals weakly keeled. elsewhere and with indistinct reddish markings near the ear and Upper labials 15-16. Femoral and preanal pores 26-31. temples. The top of the head is a dark reddish orange, The lower Distribution: C. adelynhoserae sp. nov. is known only from near flanks are characterised by 3-5 dark brown bars across a pale the type locality of the Saint George Range and nearby ranges, yellowish background, being indistinctly divided from the whitish- (e.g. Edgar Ranges) south-west Kimberley Division of Western grey above, or in some specimens these may reduce to being Australia, Australia. evenly spaced paired dark spots. There are no light blue spots or flecks anywhere on the dorsum. There is no dark bar between the Etymology: C. adelynhoserae sp. nov. is named in honour of my Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 56 Australasian Journal of Herpetology eye and ear, where the stripe would otherwise be seen in back near the mid-dorsal line is relatively indistinct and the front Ctenophorus adelynhoserae sp. nov.. legs have well defined bands. Any markings on the rear legs, if There are 17-18 tail rings, alternating blackish and whitish, the present are indistinct. rings being of even thickness. Ctenophorus adelynhoserae sp. nov. in life is depicted in Storr et Adult female C. jackyhoserae sp. nov. are of greyish colouration, al. (1983), plate 3, image 4, adult male. featuring brown barring on the lower labial area, no markings, or C. jackyhoserae sp. nov. is depicted in life in Wilson and Swan very indistinct on all limbs and a dorsal surface featuring a (2017) on page 413 at top left, adult male and Wilson and mottled appearance of light and dark, but no obvious well defined Knowles (1988), page 207, top left adult male, top right adult pattern. female and in Hoser (1989) page 67 top, male. Adult male C. lenhoseri sp. nov. have a strongly reddish-brown C. katrinahoserae sp. nov. is depicted in life online at: forebody, becoming greyish towards the rear and tail. On either https://pbase.com/gehyra/image/129646530 side of the mid-dorsal line are about 5 pairs of evenly spaced (last downloaded on 1 March 2020). semi-distinct small dark blackish brown spots of semi-rectangular C. lenhoseri sp. nov. is depicted in life in Cogger (2014), bottom shape, the longer sides being those running towards the flanks. left, adult male (aged) and Brown (2014), page 653 (3 images The flanks are more reddish than the mid dorsum, which is labelled as Ctenophorus caudicinctus macropus, Windorah, slightly greyish and the flanks also have scattered light blue spots Queensland, two adult males and an adult female). or flecks. There are 24-28 tail rings, defined by having very narrow light sections and wide darker sections, being alternating All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. brownish-black and whitish bands. There are no obvious nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. blotches or spots on the lower flanks, or dark bar between the macropus can be separated from all congeners in the subgenus eye and ear, although in some specimens there is dark peppering Tachyon Wells and Wellington, 1985 (as defined elsewhere in where the stripe would otherwise be seen in Ctenophorus this paper), by having over 32 lamellae under the fourth toe adelynhoserae sp. nov.. versus less than 32 in all other species as well as a tail 2 times the length of snout-vent, versus less than 1.8 times snout-vent in Adult female C. lenhoseri sp. nov. are of similar colour to males, all other species and the adpressed hind-leg extends past the being reddish and without obvious pattern of any sort. The dorsal snout (not so in all other species). surface is effectively unicolour, save for a poorly defined zone of darkening along the mid vertebral line and 4-6 well-spaced pairs All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. of small but obvious yellow spots on either side of the mid dorsal nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. line of the body. There are sometimes paired scattered dark spots macropus are characterised as follows: Distal three-quarters of around the neck, which typically fade in older specimens. There tail are 18-22 tail rings, with lighter ones being narrow or incomplete compressed. Nasal small, located on top of obtuse rostral ridge. and darker ones about 3-4 times wider, the colouration being Keels of dorsal scales moderately strong and sharp, terminating greyish orange (wider) and yellow (narrower) bands. Fore and in a blunt end or short spine. Ventrals weakly keeled. Upper hind limbs both have indistinct bands. labials 15-16. Femoral and preanal pores 26-31. Juvenile C. lenhoseri sp. nov. of both sexes are characterised by Distribution: C. jackyhoserae sp. nov. is known from the main a significant amount of dark brown and black pigment in the form part of the Kimberley division in Western Australia, including the of flecks or mottling on the upper surfaces, which fades with age. north and east Kimberley, as well as nearby parts of immediately Adult male C. katrinahoserae sp. nov. have an orange-grey- adjacent Northern Territory. The similar species C. adelynhoserae brown forebody, with a distinct salmon colouration across whitish sp. nov. is known only from near the type locality of the Saint parts of the upper forebody and head, becoming dull greyish George Range and nearby ranges, (e.g. Edgar Ranges) south- towards the rear and tail. On either side of the mid-dorsal line are west Kimberley Division of Western Australia, Australia. about scattered semi-distinct dark flecks which may also appear Etymology: C. jackyhoserae sp. nov. is named in honour of my on the mid flanks, but the dorsum is otherwise not prominently youngest daughter, Jacky Hoser, of Park Orchards, Victoria, marked. The flanks are slightly darker than the dorsum at the mid Australia in recognition of over 18 years active work with wildlife flanks before becoming light again at the belly. There are 20-26 conservation, research and education in Australia. tail rings, defined by having narrow light sections and wide darker CTENOPHORUS (TACHYON) KATRINAHOSERAE SP. NOV. sections, being alternating brownish-black and whitish bands. LSID urn:lsid:zoobank.org:act:8137BF13-CB27-48AB-B0CB- There are no obvious blotches or spots on the lower flanks, or F847CC8178D0 dark bar between the eye and ear, although in some specimens Holotype: A preserved specimen at the Museum and Art Gallery there is dark peppering where the stripe would otherwise be seen of the Northern Territory, Darwin, Northern Territory, Australia, in Ctenophorus adelynhoserae sp. nov.. specimen number NTM R25848 collected from 10 km south-east Adult female C. katrinahoserae sp. nov. are of similar colour to of the Roadhouse at Hells Gate, Queensland, Australia, Latitude - males, but generally greyish in colour with lighter yellow-cream 17.53 S., Longitude 138.40 E. interspaces. There is a mottled pattern across the dorsal surface, This government-owned facility allows access to its holdings. being most prominent at the anterior end and the sides of the head. Paratype: A preserved specimen at the Museum and Art Gallery of the Northern Territory, Darwin, Northern Territory, Australia, In both sexes, the legs are generally unmarked, but commonly specimen number NTM R25847 collected from 10 km south-east with indistinct blotching, particularly on the lower rear legs upper of the Roadhouse at Hells Gate, Queensland, Australia, Latitude - surface. 17.53 S., Longitude 138.40 E. Adult male C. macropus have a generally light orange dorsal Diagnosis: The species Ctenophorus katrinahoserae sp. nov. colour, with scattered dark brown flecks on the head and neck, from far north-west Queensland and the nearby parts of the but not on the body or legs. Prominent in this species are Northern Territory, south of the Gulf of Carpentaria, C. lenhoseri numerous bright aqua-blue spots scattered across the dorsal sp. nov. from the Selwyn Ranges (generally around Mount Isa surface and sides of the flanks. There are 19-21 tail rings, with and south of there), in north-west Queensland, C. jackyhoserae the darker sections being significantly wider than the lighter ones. sp. nov. from the northern and eastern Kimberley division of The anterior tail is generally brownish orange with indistinct Western Australia and immediately adjacent parts of the Northern bands, while the posterior half of the tail has fairly well defined Territory and C. adelynhoserae sp. nov. from the south-west bands, the colours being brown and yellow-white. Kimberley division of Western Australia (being the St. George Adult female C. macropus are similar in most respects to adult and Edgar Ranges, north-east of Broome, Western Australia) female C. jackyhoserae sp. nov. but any yellow spotting on the have until now all been treated as populations of C. macropus

(Storr, 1967) with a distribution centred on Arnhem Land, line of the body. There are sometimes paired scattered dark spots Northern Territory, with which they are morphologically most around the neck, which typically fade in older specimens. There similar to. are 18-22 tail rings, with lighter ones being narrow or incomplete However all are morphologically distinct, wholly allopatric and and darker ones about 3-4 times wider, the colouration being sufficiently divergent to be treated as full species as is seen greyish orange (wider) and yellow (narrower) bands. Fore and herein, with divergences estimated by Melville et al. (2016) to be hind limbs both have indistinct bands. in excess of 2.5 MYA for all but C. lenhoseri sp. nov. which was Juvenile C. lenhoseri sp. nov. of both sexes are characterised by not inspected by Melville et al. (2016). a significant amount of dark brown and black pigment in the form C. lenhoseri sp. nov. is however separated from the other of flecks or mottling on the upper surfaces, which fades with age. species by a barrier of known antiquity in far north-west Adult male C. katrinahoserae sp. nov. have an orange-grey- Queensland. brown forebody, with a distinct salmon colouration across whitish Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. nov., parts of the upper forebody and head, becoming dull greyish C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. towards the rear and tail. On either side of the mid-dorsal line are macropus are readily separated by their unique colour patterns. about scattered semi-distinct dark flecks which may also appear Adult male Ctenophorus adelynhoserae sp. nov. are separated on the mid flanks, but the dorsum is otherwise not prominently from males of the other species by having a brown dorsum and marked. The flanks are slightly darker than the dorsum at the mid flanks with either no markings or indistinct ones. Legs are either flanks before becoming light again at the belly. There are 20-26 unmarked or lack obvious markings, which otherwise may include tail rings, defined by having narrow light sections and wide darker indistinct spots or crossbands. There are scattered dark flecks or sections, being alternating brownish-black and whitish bands. peppering along the lower sides and no light blue spots or flecks There are no obvious blotches or spots on the lower flanks, or anywhere on the dorsum. Between the eye and ear is a dark and dark bar between the eye and ear, although in some specimens semi-distinct bar or broken bar. There are 19-21 tail rings, being there is dark peppering where the stripe would otherwise be seen alternating brown and whitish, the lighter sections being narrower in Ctenophorus adelynhoserae sp. nov.. than the darker sections. Adult female C. katrinahoserae sp. nov. are of similar colour to Adult female C. adelynhoserae sp. nov. are greyish in colouration males, but generally greyish in colour with lighter yellow-cream and with a greyish head with mottling as opposed to any well- interspaces. There is a mottled pattern across the dorsal surface, defined markings. being most prominent at the anterior end and the sides of the Adult male C. jackyhoserae sp. nov. normally have a brownish- head. red dorsal surface with broken lines or evenly arranged flecks In both sexes, the legs are generally unmarked, but commonly along, but not on the mid-dorsal line. Some specimens with indistinct blotching, particularly on the lower rear legs upper alternatively have a whitish-red upper body to deep red, surface. characterised by a dorsal arrangement of about six broken Adult male C. macropus have a generally light orange dorsal orange-brown-red lines running longitudinally down the body from colour, with scattered dark brown flecks on the head and neck, neck to rump, being broken by intervening areas of creamy-grey but not on the body or legs. Prominent in this species are white that are wider than the darker markings. The side of the numerous bright aqua-blue spots scattered across the dorsal head and labials are whitish or at least with whiter pigment than surface and sides of the flanks. There are 19-21 tail rings, with elsewhere and with indistinct reddish markings near the ear and the darker sections being significantly wider than the lighter ones. temples. The top of the head is a dark reddish orange, The lower The anterior tail is generally brownish orange with indistinct flanks are characterised by 3-5 dark brown bars across a pale bands, while the posterior half of the tail has fairly well defined yellowish background, being indistinctly divided from the whitish- bands, the colours being brown and yellow-white. grey above, or in some specimens these may reduce to being Adult female C. macropus are similar in most respects to adult evenly spaced paired dark spots. There are no light blue spots or female C. jackyhoserae sp. nov. but any yellow spotting on the flecks anywhere on the dorsum. There is no dark bar between the back near the mid-dorsal line is relatively indistinct and the front eye and ear, where the stripe would otherwise be seen in legs have well defined bands. Any markings on the rear legs, if Ctenophorus adelynhoserae sp. nov.. present are indistinct. There are 17-18 tail rings, alternating blackish and whitish, the Ctenophorus adelynhoserae sp. nov. in life is depicted in Storr et rings being of even thickness. al. (1983), plate 3, image 4, adult male. Adult female C. jackyhoserae sp. nov. are of greyish colouration, C. jackyhoserae sp. nov. is depicted in life in Wilson and Swan featuring brown barring on the lower labial area, no markings, or (2017) on page 413 at top left, adult male and Wilson and very indistinct on all limbs and a dorsal surface featuring a Knowles (1988), page 207, top left adult male, top right adult mottled appearance of light and dark, but no obvious well defined female and in Hoser (1989) page 67 top, male. pattern. C. katrinahoserae sp. nov. is depicted in life online at: Adult male C. lenhoseri sp. nov. have a strongly reddish-brown https://pbase.com/gehyra/image/129646530 forebody, becoming greyish towards the rear and tail. On either side of the mid-dorsal line are about 5 pairs of evenly spaced (last downloaded on 1 March 2020). semi-distinct small dark blackish brown spots of semi-rectangular C. lenhoseri sp. nov. is depicted in life in Cogger (2014), bottom shape, the longer sides being those running towards the flanks. left, adult male (aged) and Brown (2014), page 653 (3 images The flanks are more reddish than the mid dorsum, which is labelled as Ctenophorus caudicinctus macropus, Windorah, slightly greyish and the flanks also have scattered light blue spots Queensland, two adult males and an adult female). or flecks. There are 24-28 tail rings, defined by having very All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. narrow light sections and wide darker sections, being alternating nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. brownish-black and whitish bands. There are no obvious macropus can be separated from all congeners in the subgenus blotches or spots on the lower flanks, or dark bar between the Tachyon Wells and Wellington, 1985 (as defined elsewhere in eye and ear, although in some specimens there is dark peppering this paper), by having over 32 lamellae under the fourth toe where the stripe would otherwise be seen in Ctenophorus versus less than 32 in all other species as well as a tail 2 times adelynhoserae sp. nov.. the length of snout-vent, versus less than 1.8 times snout-vent in Adult female C. lenhoseri sp. nov. are of similar colour to males, all other species and the adpressed hind-leg extends past the being reddish and without obvious pattern of any sort. The dorsal snout (not so in all other species). surface is effectively unicolour, save for a poorly defined zone of All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. darkening along the mid vertebral line and 4-6 well-spaced pairs nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. of small but obvious yellow spots on either side of the mid dorsal macropus are characterised as follows: Distal three-quarters of Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 58 Australasian Journal of Herpetology tail compressed. Nasal small, located on top of obtuse rostral alternatively have a whitish-red upper body to deep red, ridge. Keels of dorsal scales moderately strong and sharp, characterised by a dorsal arrangement of about six broken terminating in a blunt end or short spine. Ventrals weakly keeled. orange-brown-red lines running longitudinally down the body from Upper labials 15-16. Femoral and preanal pores 26-31. neck to rump, being broken by intervening areas of creamy-grey Distribution: C. katrinahoserae sp. nov. is known only from hilly white that are wider than the darker markings. The side of the country south of the Gulf of Carpentaria in far northwest head and labials are whitish or at least with whiter pigment than Queensland and nearby parts of the Northern Territory, in an area elsewhere and with indistinct reddish markings near the ear and generally bounded by Hells Gate in the East and Limmen temples. The top of the head is a dark reddish orange, The lower National Park in the West. flanks are characterised by 3-5 dark brown bars across a pale C. macropus (Storr, 1967) inhabits the top end of the Northern yellowish background, being indistinctly divided from the whitish- Territory including Arnhem Land. grey above, or in some specimens these may reduce to being evenly spaced paired dark spots. There are no light blue spots or In the ranges surrounding Mount Isa and areas to the south, the flecks anywhere on the dorsum. There is no dark bar between the species C. lenhoseri sp. nov. occurs. eye and ear, where the stripe would otherwise be seen in Etymology: C. katrinahoserae sp. nov. is named in honour of my Ctenophorus adelynhoserae sp. nov.. mother, Katrina Hoser, spending most of her life in the north side There are 17-18 tail rings, alternating blackish and whitish, the of Sydney, New South Wales, Australia in recognition of over 50 rings being of even thickness. years of valuable contributions to herpetology in Australia and for Adult female C. jackyhoserae sp. nov. are of greyish colouration, services to the shoe retailing business globally. featuring brown barring on the lower labial area, no markings, or CTENOPHORUS (TACHYON) LENHOSERI SP. NOV. very indistinct on all limbs and a dorsal surface featuring a LSID urn:lsid:zoobank.org:act:524EFFEE-A159-4491-9960- mottled appearance of light and dark, but no obvious well defined DE3607193D56 pattern. Holotype: A preserved specimen at the Australian Museum, Adult male C. lenhoseri sp. nov. have a strongly reddish-brown Sydney, New South Wales, Australia, specimen number R.26001 forebody, becoming greyish towards the rear and tail. On either collected from Mount Isa in north-west Queensland, Australia, side of the mid-dorsal line are about 5 pairs of evenly spaced Latitude -20.73 S., Longitude 139.48 E. This government-owned semi-distinct small dark blackish brown spots of semi-rectangular facility allows access to its holdings. shape, the longer sides being those running towards the flanks. Paratype: A preserved specimen at the Australian Museum, The flanks are more reddish than the mid dorsum, which is Sydney, New South Wales, Australia, specimen number R.72760 slightly greyish and the flanks also have scattered light blue spots collected from Mount Isa in north-west Queensland, Australia, or flecks. There are 24-28 tail rings, defined by having very Latitude -20.73 S., Longitude 139.48 E. narrow light sections and wide darker sections, being alternating Diagnosis: The species Ctenophorus lenhoseri sp. nov. from the brownish-black and whitish bands. There are no obvious Selwyn Ranges (generally around Mount Isa and south of there), blotches or spots on the lower flanks, or dark bar between the in north-west Queensland, C. katrinahoserae sp. nov. from far eye and ear, although in some specimens there is dark peppering north-west Queensland and the nearby parts of the Northern where the stripe would otherwise be seen in Ctenophorus Territory, south of the Gulf of Carpentaria, C. jackyhoserae sp. adelynhoserae sp. nov.. nov. from the northern and eastern Kimberley division of Western Adult female C. lenhoseri sp. nov. are of similar colour to males, Australia and immediately adjacent parts of the Northern Territory being reddish and without obvious pattern of any sort. The dorsal and C. adelynhoserae sp. nov. from the south-west Kimberley surface is effectively unicolour, save for a poorly defined zone of division of Western Australia (being the St. George and Edgar darkening along the mid vertebral line and 4-6 well-spaced pairs Ranges, north-east of Broome, Western Australia) have until now of small but obvious yellow spots on either side of the mid dorsal all been treated as populations of C. macropus (Storr, 1967) with line of the body. There are sometimes paired scattered dark spots a distribution centred on Arnhem Land, Northern Territory, with around the neck, which typically fade in older specimens. There which they are morphologically most similar to. are 18-22 tail rings, with lighter ones being narrow or incomplete However all are morphologically distinct, wholly allopatric and and darker ones about 3-4 times wider, the colouration being sufficiently divergent to be treated as full species as is seen greyish orange (wider) and yellow (narrower) bands. Fore and herein, with divergences estimated by Melville et al. (2016) to be hind limbs both have indistinct bands. in excess of 2.5 MYA for all but C. lenhoseri sp. nov. which was Juvenile C. lenhoseri sp. nov. of both sexes are characterised by not inspected by Melville et al. (2016). a significant amount of dark brown and black pigment in the form C. lenhoseri sp. nov. is however separated from the other of flecks or mottling on the upper surfaces, which fades with age. species by a barrier of known antiquity in far north-west Adult male C. katrinahoserae sp. nov. have an orange-grey- Queensland. brown forebody, with a distinct salmon colouration across whitish Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. nov., parts of the upper forebody and head, becoming dull greyish C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. towards the rear and tail. On either side of the mid-dorsal line are macropus are readily separated by their unique colour patterns. about scattered semi-distinct dark flecks which may also appear Adult male Ctenophorus adelynhoserae sp. nov. are separated on the mid flanks, but the dorsum is otherwise not prominently from males of the other species by having a brown dorsum and marked. The flanks are slightly darker than the dorsum at the mid flanks with either no markings or indistinct ones. Legs are either flanks before becoming light again at the belly. There are 20-26 unmarked or lack obvious markings, which otherwise may include tail rings, defined by having narrow light sections and wide darker indistinct spots or crossbands. There are scattered dark flecks or sections, being alternating brownish-black and whitish bands. peppering along the lower sides and no light blue spots or flecks There are no obvious blotches or spots on the lower flanks, or anywhere on the dorsum. Between the eye and ear is a dark and dark bar between the eye and ear, although in some specimens semi-distinct bar or broken bar. There are 19-21 tail rings, being there is dark peppering where the stripe would otherwise be seen alternating brown and whitish, the lighter sections being narrower in Ctenophorus adelynhoserae sp. nov.. than the darker sections. Adult female C. katrinahoserae sp. nov. are of similar colour to Adult female C. adelynhoserae sp. nov. are greyish in colouration males, but generally greyish in colour with lighter yellow-cream and with a greyish head with mottling as opposed to any well- interspaces. There is a mottled pattern across the dorsal surface, defined markings. being most prominent at the anterior end and the sides of the Adult male C. jackyhoserae sp. nov. normally have a brownish- head. red dorsal surface with broken lines or evenly arranged flecks In both sexes, the legs are generally unmarked, but commonly along, but not on the mid-dorsal line. Some specimens with indistinct blotching, particularly on the lower rear legs upper Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 59

surface. allows access to its holdings. Adult male C. macropus have a generally light orange dorsal Paratype: A preserved specimen at the Museum and Art Gallery colour, with scattered dark brown flecks on the head and neck, of the Northern Territory, Reptile Collection, Darwin, Northern but not on the body or legs. Prominent in this species are Territory, Australia, specimen number R31309 collected from numerous bright aqua-blue spots scattered across the dorsal Devils Marbles, Northern Territory, Australia, Latitude -20.55, surface and sides of the flanks. There are 19-21 tail rings, with Longitude 134.283 E. the darker sections being significantly wider than the lighter ones. Diagnosis: Until now Ctenophorus maxinehoserae sp. nov. has The anterior tail is generally brownish orange with indistinct been treated as a northern population of C. slateri (Storr, 1967), bands, while the posterior half of the tail has fairly well defined the latter species with a type locality of Hermannsburg, Northern bands, the colours being brown and yellow-white. Territory, Australia, Latitude 23.58 S., Longitude 132.46 E. Adult female C. macropus are similar in most respects to adult C. slateri as defined herein is effectively confined to the female C. jackyhoserae sp. nov. but any yellow spotting on the Macdonnell Ranges of Central Australia. back near the mid-dorsal line is relatively indistinct and the front However genetic divergence and morphological differences, legs have well defined bands. Any markings on the rear legs, if combined with a clearly allopatric population indicate that the present are indistinct. lizards herein identified as Ctenophorus maxinehoserae sp. nov. Ctenophorus adelynhoserae sp. nov. in life is depicted in Storr et found to the north of the type region of C. slateri are not of the al. (1983), plate 3, image 4, adult male. same species as C. slateri (Storr, 1967). C. jackyhoserae sp. nov. is depicted in life in Wilson and Swan Both C. maxinehoserae sp. nov. and C. slateri are readily (2017) on page 413 at top left, adult male and Wilson and separated from all other species in the subgenus Tachyon (Wells Knowles (1988), page 207, top left adult male, top right adult and Wellington, 1985) (as defined elsewhere in this paper) by the female and in Hoser (1989) page 67 top, male. following unique suite of characters: Keels of dorsal scales sharp C. katrinahoserae sp. nov. is depicted in life online at: and black (versus not so in all other species), the adpressed https://pbase.com/gehyra/image/129646530 hind-leg does not extend to the end of the snout, which clearly separates this species from all of Ctenophorus adelynhoserae (last downloaded on 1 March 2020). sp. nov., C. jackyhoserae sp. nov., C. katrinahoserae sp. nov., C. C. lenhoseri sp. nov. is depicted in life in Cogger (2014), bottom lenhoseri sp. nov. and C. macropus (Storr, 1967) (these species left, adult male (aged) and Brown (2014), page 653 (3 images all previously treated as populations of C. macropus), base of tail labelled as Ctenophorus caudicinctus macropus, Windorah, not compressed; there are no dark blackish caudal bands, nasal Queensland, two adult males and an adult female). on or just below a swollen rostral ridge: pores fewer than 43, All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. colouration with a dorsal surface that is reddish-brown or dull nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. orange, with small indistinct, whitish or darkish specks, spots or macropus can be separated from all congeners in the subgenus small blotches. Tail rings are semidistinct at the anterior end and Tachyon Wells and Wellington, 1985 (as defined elsewhere in fully formed at the rear, with darker sections 3-5 times wider, and this paper), by having over 32 lamellae under the fourth toe a colouration of alternating greyish orange and yellowish white versus less than 32 in all other species as well as a tail 2 times bands. the length of snout-vent, versus less than 1.8 times snout-vent in Both male and female C. maxinehoserae sp. nov. are readily all other species and the adpressed hind-leg extends past the separated from C. slateri by having a series of 6-7 small semi- snout (not so in all other species). distinct dark purplish-brown spots running along either side of the All of Ctenophorus adelynhoserae sp. nov., C. jackyhoserae sp. mid dorsal line of the body with further similarly-spaced similar nov., C. katrinahoserae sp. nov., C. lenhoseri sp. nov. and C. markings extending to the upper surface of the base of the tail. macropus are characterised as follows: Distal three-quarters of Both male and female C. slateri are characterised by having a tail compressed. Nasal small, located on top of obtuse rostral series of well-defined white to whitish yellow spots running along ridge. Keels of dorsal scales moderately strong and sharp, either side of the mid dorsal line of the body with further similarly- terminating in a blunt end or short spine. Ventrals weakly keeled. spaced similar but less distinct markings extending to the upper Upper labials 15-16. Femoral and preanal pores 26-31. surface of the base of the tail. Distribution: The species C. lenhoseri sp. nov. occurs in the C. maxinehoserae sp. nov. is further separated from C. slateri by ranges surrounding Mount Isa and areas to the south in north- having weakly keeled ventral scales, versus not so in C. slateri. west Queensland. C. maxinehoserae sp. nov. in life is depicted in Wilson and C. katrinahoserae sp. nov. is known only from hilly country south Knowles (1988), on page 206 bottom right, adult male, and of the Gulf of Carpentaria in far northwest Queensland and Brown (2014), page 653, bottom left. nearby parts of the Northern Territory, in an area generally C. slateri in life is depicted in Brown (2014) on page 653 bottom bounded by Hells Gate in the East and Limmen National Park in right, adult female. the West. Distribution: C. maxinehoserae sp. nov. is restricted to ranges C. macropus (Storr, 1967) inhabits the top end of the Northern and rocky country in the general vicinity of Barrow Creek to Territory including Arnhem Land. Tennant Creek, including the Davenport Ranges in the Northern Etymology: C. lenhoseri sp. nov. is named in honour of my Territory, Australia, while C. slateri is confined to the Macdonnell father, Len Hoser (AKA Leonard Donald Hoser), spending about Ranges and nearby rocky hills in Central Australia. half of his life living in the north side of Sydney, New South Etymology: Named in honour of Maxine Hoser of Margate in the Wales, Australia and the other half in the United Kingdom of United Kingdom of England, Wales, Scotland and Northern England, Wales, Scotland and Northern Ireland in recognition of Ireland in recognition of her services to herpetology from the over three decades of valuable contributions to herpetology in 1960’s to 1980’s. Australia and for services to the bakery business in Australia and CTENOPHORUS (TACHYON) RONHOSERI SP. NOV. the UK. LSID urn:lsid:zoobank.org:act:6BA8B610-14A6-40B6-A681- CTENOPHORUS (TACHYON) MAXINEHOSERAE SP. NOV. E7F0AD96F3A8 LSID urn:lsid:zoobank.org:act:EB5ADF71-70FB-45A9-B989- Holotype: A preserved specimen at the Western Australian 5C107063D718 Museum, Perth, Western Australia, Australia, specimen number Holotype: A preserved specimen at the National Museum of R102084, collected from the Yulpul Rockhole at the north end of Victoria, Melbourne, Victoria, Australia, specimen number the McKay Range, Western Australia, Australia, Latitude -22.97 D74268 collected from Devils Marbles, Northern Territory, S., Longitude 122.46 E. This government-owned facility allows Australia, Latitude -20.5676, Longitude 134.264 E. This facility access to its holdings. Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved 60 Australasian Journal of Herpetology

Paratype: A preserved specimen at the Western Australian shireenhoserae sp. nov. and C. ornatus by colouration. In this Museum, Perth, Western Australia, Australia, specimen number species the mid dorsal blotches are similar in form to those seen R102611, collected from the Little Sandy Desert, Western in C. ornatus including being bounded by black. But unlike C. Australia, Australia, Latitude -24.05 S., Longitude 120.41 E. ornatus that has a generally black dorsum and flanks with some Diagnosis: Morphologically, Ctenophorus ronhoseri sp. nov. is obvious yellow spots, C. sharonhoserae sp. nov. has a reddish most similar to C. caudicinctus (Günther, 1875), although brown dorsum and flanks, with a distinctive line boundary genetically it is most closely related to C. graafi (Storr, 1967). between the reddish brown dorsum and white undersides. At However the species is sufficiently divergent from all congeners most there are only a small number of very tiny whitish yellow (4.2 MYA from nearest common ancestor according to the specks on the lower flanks and the mid flanks have no such evidence of Melville et al. 2016), morphologically distinct and markings, being either dark or light reddish-brown in colour. reproductively isolated as to warrant being recognized as a full In C. sharonhoserae sp. nov. the forelimbs are well-banded and species. the hind limbs are not, which places this taxon intermediate Ctenophorus ronhoseri sp. nov. and C. caudicinctus are between C. shireenhoserae sp. nov. whose limbs lack obvious separated from all other species in the subgenus Tachyon (Wells distinct bands and C. ornatus which has well banded front and and Wellington) as defined elsewhere in this paper, including the hind limbs. closely related species C. graafi (Storr, 1967) by having the nasal The species C. shireenhoserae sp. nov. is in many ways on or just below a swollen rostral ridge: pores fewer than 43 and intermediate in form between C. ornatus and C. yinnietherra the whole of tail compressed with dark narrow caudal bands (Storr, 1981). C. yinnietherra is separated from C. encircling the tail. shireenhoserae sp. nov., C. sharonhoserae sp. nov. and C. C. ronhoseri sp. nov. is readily separated from C. caudicinctus by ornatus by having the basal portion of the tail unbanded and only having yellowish or orangeish colouration on the back, versus banding on the posterior third. C. yinnietherra is further reddish or brown on C. caudicinctus. Markings on the back and separated from both C. shireenhoserae sp. nov., C. flanks of adult male C. ronhoseri sp. nov. are indistinct, versus sharonhoserae sp. nov. and C. ornatus by dorsal colouration. distinct in C. caudicinctus. The flanks are generally greyish, without obvious markings, forelimbs also greyish and mid-dorsal area and rear limbs C. caudicinctus also has white marks or peppering on the sides generally reddish-orange and without any obvious black markings of the head, whereas this is not the case in C. ronhoseri sp. nov.. or black bounding any blotches or spots (which are absent). The Distribution: C. ronhoseri sp. nov. is known only from the type anterior half of the tail has an orange upper surface (without localities and immediately adjacent hilly areas in the east Pilbara bands), versus not orangeish and banded in the other three region of Western Australia, Australia. species. C. caudicinctus is found in other parts of the Pilbara, including C. yinnietherra is further separated from C. shireenhoserae sp. the west Pilbara nov., C. sharonhoserae sp. nov. and C. ornatus by having a Etymology: Named in honour of Ron Hoser, (deceased) of significant amount of orange around the eye and nearby parts of Green Valley, (Sydney), New South Wales, Australia in the head (absent in the other two species). recognition of numerous contributions to herpetology in the Adult female C. yinnietherra have a generally unbanded tail, 1960’s to 1980’s. versus banded in females of C. shireenhoserae sp. nov., C. CTENOPHORUS (TACHYON) SHARONHOSERAE SP. NOV. sharonhoserae sp. nov. and C. ornatus. Adult female C. LSID urn:lsid:zoobank.org:act:2FC73FDD-6F04-41F3-91D6- shireenhoserae sp. nov. and C. sharonhoserae sp. nov. are 3676BB32C04B generally a reddish-brown dorsally with semi-distinct irregular Holotype: A preserved specimen at the Western Australian blackish markings, versus greyish-brown dorsally with semi- Museum, Perth, Western Australia, Australia, specimen number: distinct irregular blackish markings in C. ornatus. R12734, collected at Dedari, (Coolgardie), Western Australia, C. shireenhoserae sp. nov. in life is depicted in Cogger (2014) on Australia, Latitude -31.08 S., Longitude 120.77 E. This page 716 at top right; Storr, Smith and Johnstone (1983) middle government-owned facility allows access to its holdings. right and in Brown (2014) on page 656 at bottom left (with caption Paratypes: Two preserved specimens at the Western Australian transposed from photo of C. ornatus in middle right of same Museum, Perth, Western Australia, Australia, specimen numbers page). R29946 and R29947 collected at Widgiemooltha, Western C. ornatus in life is depicted in Cogger (2014) on page 717 at top Australia, Australia, Latitude -31.3 S., Longitude 121.58 E. left; Storr, Smith and Johnstone (1983) top right and left and in Diagnosis: Until now C. sharonhoserae sp. nov. has been Brown (2014) on page 656 at middle right (with caption treated as an inland eastern population of C. ornatus (Gray, transposed from photo of shireenhoserae sp. nov. in bottom left 1845) with a distribution mainly in the Goldfields region of of same page). southern Western Australia, Australia. C. sharonhoserae sp. nov. in life is depicted in Storr, Smith and These two species and third species formally described herein as Johnstone (1983) middle left. C. shireenhoserae sp. nov. are readily separated by colouration C. yinnietherra in life is depicted in Cogger (2014) on page 724 at in adult males. top and Storr, Smith and Johnstone (1983), bottom right and left. The type form of C. ornatus has a generally black dorsal surface The type specimen of C. ornatus as described by Boulenger with large creamy-white blotches running down the mid-dorsal (1885), as described by him in terms of colouration, clearly line and a lesser number of smaller spots on the flanks. The tail conforms to a male specimen from the wetter parts of South- has well defined black and yellow-white bands (sometimes western Western Australia (as in somewhere near Perth) and so uneven) of similar thickness on the dorsal surface and blackish that form is regarded as nominate for C. ornatus, thereby limbs that are strongly banded or marked with yellow-white. confirming the taxon herein described as C. shireenhoserae sp. By contrast C. shireenhoserae sp. nov. has a generally reddish- nov. was previously undescribed. brown dorsal surface with an absence of yellow spots on the side. In terms of colouration, Boulenger wrote: “Black above; a series The colour of the mid dorsal blotches is whitish-grey, generally of large irregular yellowish spots along the vertebral line and a merging to form a mid-dorsal stripe and bounded by dark brown few very small ones scattered on the sides; limbs and tail with to black. The limbs are generally a light greyish colour and with ill yellowish cross bars; throat punctate with blackish; a large black defined blotches or markings. The tail of C. shireenhoserae sp. spot covers the chest.” nov. is banded but the lighter interspaces are double the size of Boulenger’s full description of the type specimen for the darker sections. The lighter bands are light bluish-grey, Gray’s C. ornatus was as follows: versus white to fellow-white in C. ornatus. “ H a bit slender . Head moder ately large; snout slightl y long er

C. sharonhoserae sp. nov. is separated from both C. Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved Australasian Journal of Herpetology 61

than the diameter of the orbit; canthus r ostralis swollen, not femoral and preanal pores present in males; adult males usually angular ; nostr il distinctl y tub ular, directed slightl y upwards, with distinctive black or dark grey markings on the throat and/or muc h near er the e ye than the end of the snout ; tympan u m chest. large, three f ifths the diameter of the orbit ; upper head- The genus Paractenophorus Hoser, 2015 is separated from scales tuber cular, r ough, smallest on supr aorbital region. Ctenophorus, Notactenophorus Hoser, 2015 and Sides of neck rather strongly plicate a distinct dorso-lateral Pseudoctenophorus Hoser, 2015 by the following suite of fold. Gular scales minute, smooth. Body much depressed, characters: tympanum exposed; no series of enlarged, spinose covered a bo ve with small k eeled scales, lar gest and unif orm scales on either side of the base of the tail; hind limb reaching no on the v ertebral region, min ute and inter mix ed with widel y further than the tympanum when adpressed; tail usually less than scattered slightl y enlar g ed ones on the sides; no dor sal crest; 1.5 times as long as the head and body; nasal region is not ventr al scales small, smooth. Limbs and dig its long, the swollen, the nostril lying below an angular canthal ridge; pores adpressed hind limb reaching the tip of the snout; the scales fewer than 15; nostril is slit-like or narrowly elliptical. on the limbs str ongl y k eeled , those on the ar m and tibia Specimens within the genus Pseudoctenophorus Hoser, 2015 muc h enlar g ed. are separated from all other Ctenophorus Fitzinger, 1843, the A ser ies of sixty por es e xtending along the w hole length of genus they were placed previously, by the following suite of the thighs, slightly interrupted on the pr aeanal r eg ion. Tail characters, being one or other of the following three: slender , r ound , depr essed a t the base; twice as long as head 1/ Tympanum exposed; a series of enlarged, spinose scales on and body; either side of the base of the tail (subgenus Pseudoctenophorus caudal scales equal, m uch lar g er than dor sals, str ongl y Hoser, 2015), or: keeled . 2/ Tympanum hidden; covered by skin; body scales are strongly Blac k above; a series of large irregular y ello wish spots heterogeneous, many of the larger scales on the body and head along the v ertebral line and a few ver y small ones sca ttered keeled or moderately spinose; a series of enlarged, spinose on the sides; limbs and tail with y ello wish cr oss bar s; throat scales on either side of the base of the tail (subgenus punctate with Chapmanagama Hoser, 2015), or: blackish; a large blac k spot co vers the chest.” 3/ Tympanum hidden; covered by skin; body scales are strongly Species within the subgenus Tachyon Wells and Wellington, heterogeneous, many of the larger scales on the body and head 1985 are separated from all other Ctenophorus Fitzinger, 1843 by keeled or moderately spinose; no series of enlarged, spinose the following suite of characters being one or other of: scales on either side of the base of the tail (subgenus 1/ Tympanum exposed; no series of enlarged, spinose scales on Turnbullagama Hoser, 2015). either side of the base of the tail; hind limb usually reaching to Notactenophorus Hoser, 2015 is readily separated from all other eye or beyond when adpressed; tail usually much more than 1.5 members of the genus Ctenophorus (where it has been placed times as long as the head and body; canthus rostralis swollen, until now, as defined in Cogger 2014) and Pseudoctenophorus but nostrils, when viewed from above, face distinctly upwards as Hoser, 2015 by the following unique suite of characters: opposed to outwards (species C. adelynhoserae sp. nov.; C. Tympanum is hidden being covered by skin, the body scales are caudicinctus (Günther, 1875); C. graafi (Storr, 1967); C. smooth, mostly small, homogenous, with scattered larger but imbricatus (Peters, 1876); C. infans (Storr, 1967); C. small, flat scales, not keeled or spinose, with a dorsal pattern of a jackyhoserae sp. nov.; C. katrinahoserae sp. nov.; C. lenhoseri longitudinal dorso-lateral series of five or six large black spots on sp. nov.; C. macropus (Storr, 1967); C. mensarum (Storr, 1967); either side. C. slateri (Storr, 1967)) or: Distribution: C. sharonhoserae sp. nov. is found in the semi-arid 2/ Tympanum exposed; no series of enlarged, spinose scales on zone of south-western Western Australia, away from the Darling either side of the base of the tail; hind limb usually reaching to Range and south coast of Western Australia, and also south of eye or beyond when adpressed; tail usually much more than 1.5 the mulga-eucalypt line in south-west Australia. The distribution is times as long as the head and body; canthus rostralis angular or mainly in the Goldfields region of southern Western Australia, moderately swollen, but nostrils, when viewed from above, face Australia. outwards as opposed to distinctly upwards (as seen in the C ornatus is restricted to the Darling Range (near Perth) and species C. adelynhoserae sp. nov.; C. caudicinctus (Günther, wetter parts of the south coast of south-western Australia. 1875); C. graafi (Storr, 1967); C. imbricatus (Peters, 1876); C. C. shireenhoserae sp. nov. is known only from near the type infans (Storr, 1967); C. jackyhoserae sp. nov.; C. katrinahoserae locality and nearby areas in Western Australia, generally between sp. nov.; C. lenhoseri sp. nov.; C. macropus (Storr, 1967); C. Paynes Find and Mount Magnet in Western Australia. mensarum (Storr, 1967); C. slateri (Storr, 1967)); at most a few Etymology: C. sharonhoserae sp. nov. is named in honour of my enlarged keeled scales on the nape; a series of enlarged cousin Sharon Hoser (now Menzies), originally of the UK, but vertebral scales, if present, forming a distinct linear series only to since having resided in various capital cities of Australia and also about the level of the forelimbs; dorsal scales at most with low, in Papua New Guinea in recognition of her contributions to irregular keels which do not form distinct continuous ridges; herpetology in the 1960’s and 1970’s. dorsolateral scales and those on the chest smooth, or with low CTENOPHORUS (TACHYON) SHIREENHOSERAE SP. NOV. blunt edges; nostril elliptical in a swollen nasal scale lying on a swollen canthal ridge; tibial region with a series of anterior LSID urn:lsid:zoobank.org:act:D6B994AC-026E-4285-9F46- proximal scales which are very much larger than those on the 51CB695A73BB posterior surface (species C. shireenhoserae sp. nov.; C. ornatus Holotype: A preserved specimen at the Western Australian and C. yinnietharra). Museum, Perth, Western Australia, Australia, specimen number: Ctenophorus as defined until now (Cogger 2014, Hoser 2015g) is R117278 collected 7 km north of Mount Magnet, Western defined by the following definition, modified to take into account Australia, Latitude -28 S., Longitude 117.87 E. This government- the new genera as defined herein. Ctenophorus is defined as an owned facility allows access to its holdings. Australian agamid genus characterised by small dorsal scales, Paratype: A preserved specimen at the Western Australian homogenous or with at most slightly enlarged tubercles; a few Museum, Perth, Western Australia, Australia, specimen number: species with distinct rows of paravertebral or dorsolateral spinose R117279 collected 7 km north of Mount Magnet, Western scales; a row of enlarged scales from below the eye to above the Australia, Latitude -28 S., Longitude 117.87 E. ear; tympanum exposed (not exposed in Notactenophorus Hoser, Diagnosis: Until now C. shireenhoserae sp. nov. has been 2015 and most Pseudoctenophorus Hoser, 2015); tail long, treated as a north-west population of C. ornatus (Gray, 1845). ranging from slightly to much longer than the head and body; These two species and third species formally described herein as

C. sharonhoserae sp. nov. are readily separated by colouration in Johnstone (1983) middle left. adult males. C. yinnietherra in life is depicted in Cogger (2014) on page 724 at The type form of C. ornatus has a generally black dorsal surface top and Storr, Smith and Johnstone (1983), bottom right and left. with large creamy-white blotches running down the mid-dorsal The type specimen of C. ornatus as described by Boulenger line and a lesser number of smaller spots on the flanks. The tail (1885), as described by him in terms of colouration, clearly has well defined black and yellow-white bands (sometimes conforms to a male specimen from the wetter parts of South- uneven) of similar thickness on the dorsal surface and blackish western Western Australia (as in somewhere near Perth) and so limbs that are strongly banded or marked with yellow-white. that form is regarded as nominate for C. ornatus, thereby By contrast C. shireenhoserae sp. nov. has a generally reddish- confirming the taxon herein described as C. shireenhoserae sp. brown dorsal surface with an absence of yellow spots on the side. nov. was previously undescribed. The colour of the mid dorsal blotches is whitish-grey, generally In terms of colouration, Boulenger wrote: “Black above; a series merging to form a mid-dorsal stripe and bounded by dark brown of large irregular yellowish spots along the vertebral line and a to black. The limbs are generally a light greyish colour and with ill few very small ones scattered on the sides; limbs and tail with defined blotches or markings. The tail of C. shireenhoserae sp. yellowish cross bars; throat punctate with blackish; a large black nov. is banded but the lighter interspaces are double the size of spot covers the chest.” the darker sections. The lighter bands are light bluish-grey, Boulenger’s full description of the type specimen for Gray’s C. versus white to fellow-white in C. ornatus. ornatus was as follows: C. sharonhoserae sp. nov. is separated from both C. “Habit slender. Head moderately large; snout slightly longer than shireenhoserae sp. nov. and C. ornatus by colouration. In this the diameter of the orbit; canthus rostralis swollen, not angular ; species the mid dorsal blotches are similar in form to those seen nostril distinctly tubular, directed slightly upwards, much nearer in C. ornatus including being bounded by black. But unlike C. the eye than the end of the snout ; tympanum large, three fifths ornatus that has a generally black dorsum and flanks with some the diameter of the orbit ; upper head-scales tubercular, rough, obvious yellow spots, C. sharonhoserae sp. nov. has a reddish smallest on supraorbital region. Sides of neck rather strongly brown dorsum and flanks, with a distinctive line boundary plicate a distinct dorso-lateral fold. Gular scales minute, smooth. between the reddish brown dorsum and white undersides. At Body much depressed, covered above with small keeled scales, most there are only a small number of very tiny whitish yellow largest and uniform on the vertebral region, minute and specks on the lower flanks and the mid flanks have no such intermixed with widely scattered slightly enlarged ones on the markings, being either dark or light reddish-brown in colour. sides; no dorsal crest; ventral scales small, smooth. Limbs and In C. sharonhoserae sp. nov. the forelimbs are well-banded and digits long, the adpressed hind limb reaching the tip of the snout; the hind limbs are not, which places this taxon intermediate the scales on the limbs strongly keeled, those on the arm and between C. shireenhoserae sp. nov. whose limbs lack obvious tibia much enlarged. distinct bands and C. ornatus which has well banded front and A series of sixty pores extending along the whole length of the hind limbs.. thighs, slightly interrupted on the praeanal region. Tail slender, The species C. shireenhoserae sp. nov. is in many ways round, depressed at the base; twice as long as head and body; intermediate in form between C. ornatus and C. yinnietherra caudal scales equal, much larger than dorsals, strongly keeled. (Storr, 1981). C. yinnietherra is separated from C. Black above; a series of large irregular yellowish spots along the shireenhoserae sp. nov., C. sharonhoserae sp. nov. and C. vertebral line and a few very small ones scattered on the sides; ornatus by having the basal portion of the tail unbanded and only limbs and tail with yellowish cross bars; throat punctate with banding on the posterior third. C. yinnietherra is further separated from both C. shireenhoserae sp. nov., C. blackish; a large black spot covers the chest.” sharonhoserae sp. nov. and C. ornatus by dorsal colouration. Species within the subgenus Tachyon Wells and Wellington, The flanks are generally greyish, without obvious markings, 1985 are separated from all other Ctenophorus Fitzinger, 1843 by forelimbs also greyish and mid-dorsal area and rear limbs the following suite of characters being one or other of: generally reddish-orange and without any obvious black markings 1/ Tympanum exposed; no series of enlarged, spinose scales on or black bounding any blotches or spots (which are absent). The either side of the base of the tail; hind limb usually reaching to anterior half of the tail has an orange upper surface (without eye or beyond when adpressed; tail usually much more than 1.5 bands), versus not orangeish and banded in the other three times as species. long as the head and body; canthus rostralis swollen, but nostrils, C. yinnietherra is further separated from C. shireenhoserae sp. when viewed from above, face distinctly upwards as opposed to nov., C. sharonhoserae sp. nov. and C. ornatus by having a outwards (species C. adelynhoserae sp. nov.; C. caudicinctus significant amount of orange around the eye and nearby parts of (Günther, 1875); C. graafi (Storr, 1967); C. imbricatus (Peters, the head (absent in the other two species). 1876); C. infans (Storr, 1967); C. jackyhoserae sp. nov.; C. Adult female C. yinnietherra have a generally unbanded tail, katrinahoserae sp. nov.; C. lenhoseri sp. nov.; C. macropus versus banded in females of C. shireenhoserae sp. nov., C. (Storr, 1967); C. mensarum (Storr, 1967); C. slateri (Storr, 1967)) sharonhoserae sp. nov. and C. ornatus. Adult female C. or: shireenhoserae sp. nov. and C. sharonhoserae sp. nov. are 2/ Tympanum exposed; no series of enlarged, spinose scales on generally a reddish-brown dorsally with semi-distinct irregular either side of the base of the tail; hind limb usually reaching to blackish markings, versus greyish-brown dorsally with semi- eye or beyond when adpressed; tail usually much more than 1.5 distinct irregular blackish markings in C. ornatus. times as long as the head and body; canthus rostralis angular or C. shireenhoserae sp. nov. in life is depicted in Cogger (2014) on moderately swollen, but nostrils, when viewed from above, face page 716 at top right; Storr, Smith and Johnstone (1983) middle outwards as opposed to distinctly upwards (as seen in the right and in Brown (2014) on page 656 at bottom left (with caption species C. adelynhoserae sp. nov.; C. caudicinctus (Günther, transposed from photo of C. ornatus in middle right of same 1875); C. graafi (Storr, 1967); C. imbricatus (Peters, 1876); C. page). infans (Storr, 1967); C. jackyhoserae sp. nov.; C. katrinahoserae C. ornatus in life is depicted in Cogger (2014) on page 717 at top sp. nov.; C. lenhoseri sp. nov.; C. macropus (Storr, 1967); C. left; Storr, Smith and Johnstone (1983) top right and left and in mensarum (Storr, 1967); C. slateri (Storr, 1967)); Brown (2014) on page 656 at middle right (with caption at most a few enlarged keeled scales on the nape; a series of transposed from photo of shireenhoserae sp. nov. in bottom left enlarged vertebral scales, if present, forming a distinct linear of same page). series only to about the level of the forelimbs; dorsal scales at C. sharonhoserae sp. nov. in life is depicted in Storr, Smith and most with low,

irregular keels which do not form distinct continuous ridges; REFERENCES CITED dorsolateral scales and those on the chest smooth, or with low Ahl, E. 1926. Neue Eidechsen und Amphibien. Zool. Anz. 67:186- blunt edges; nostril elliptical in a swollen nasal scale lying on a 192. swollen canthal ridge; tibial region with a series of anterior Baverstock, P. R. and Bradshaw, S. D. 1975. Variation in rate of proximal scales which are very much larger than those on the growth and adrenal corticosteroidogenesis in field and laboratory posterior surface (species C. shireenhoserae sp. nov.; C. ornatus populations of the lizard Amphibolurus ornatus. Comp Biochem and C. yinnietharra). Physiol 52A:557-566. Ctenophorus as defined until now (Cogger 2014, Hoser 2015g) is Boulenger, G. A. 1885. Catalogue of the lizards in the British defined by the following definition, modified to take into account Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplatidae, the new genera as defined herein. Ctenophorus is defined as an Pygopodidae, Agamidae. London:UK:450 pp. Australian agamid genus characterised by small dorsal scales, Bradshaw, S. D. 1970. Seasonal changes in the water and homogenous or with at most slightly enlarged tubercles; a few electrolyte metabolism of Amphibolurus lizards in the field. Comp species with distinct rows of paravertebral or dorsolateral spinose Biochem Physiol 36:689-718. scales; a row of enlarged scales from below the eye to above the Bradshaw, S. D. 1971. Growth and mortality in a field population ear; tympanum exposed (not exposed in Notactenophorus Hoser, of Amphibolurus lizards exposed to seasonal cold and aridity. 2015 and most Pseudoctenophorus Hoser, 2015); tail long, Journal of Zoology 165:1-25. ranging from slightly to much longer than the head and body; femoral and preanal pores present in males; adult males usually Bradshaw, S. D. and Main, A. R. 1968. Behavioural attitudes and with obvious black or dark grey markings on throat and/or chest. regulation of temperature in Amphibolurus lizards. Journal of Zoology 154:193-221. The genus Paractenophorus Hoser, 2015 is separated from Ctenophorus, Notactenophorus Hoser, 2015 and Bradshaw, S. D. and Shoemaker, V. H. 1967. Aspects of water Pseudoctenophorus Hoser, 2015 by the following suite of and electrolyte changes in a field population of Amphibolurus characters: tympanum exposed; no series of enlarged, spinose lizards. Comp Biochem Physiol 20:855-865. scales on either side of the base of the tail; hind limb reaching no Cogger, H. G. 2014. Reptiles and Amphibians of Australia, 7th further than the tympanum when adpressed; tail usually less than ed. CSIRO Publishing, Australia:xxx+1033 pp. 1.5 times as long as the head and body; nasal region is not Cogger, H. G., Cameron, E. E. and Cogger, H. M. 1983. swollen, the nostril lying below an angular canthal ridge; pores Zoological Catalogue of Australia (1) Amphibia and Reptilia. fewer than 15; nostril is slit-like or narrowly elliptical. Aust. Gov. Publishing Service, Canberra, ACT, Australia:319 pp. Specimens within the genus Pseudoctenophorus Hoser, 2015 Court of Appeal Victoria. 2014. Hoser v Department of are separated from all other Ctenophorus Fitzinger, 1843, the Sustainability and Environment [2014] VSCA 206 (5 Sept. 2014). genus they were placed previously, by the following suite of Denzer, W., Günther, R. and Manthey, U. 1997. Annotated type characters, being one or other of the following three: catalogue of the Agamid lizards (Reptilia: Squamata: Agamidae) 1/ Tympanum exposed; a series of enlarged, spinose scales on in the Museum für Naturkunde der Humbolt-Universität zu Berlin either side of the base of the tail (subgenus Pseudoctenophorus (former Zoological Museum Berlin). Mitt. Zool. Mus. Berlin Hoser, 2015), or: 73(2):309-332. 2/ Tympanum hidden; covered by skin; body scales are strongly Doody, J. S. and Schembri, B. 2014. Ctenophorus caudicinctus heterogeneous, many of the larger scales on the body and head (ring-tailed dragon) predation. Herp. Review 45(3):494-495. keeled or moderately spinose; a series of enlarged, spinose Dubois, A., Bauer, A. M., Ceriaco, L. M. P., Dusoulier, F., Fretey, scales on either side of the base of the tail (subgenus T., Lobl, I., Lorvelec, O., Ohler, A., Stopiglia, R. and Aescht, E. Chapmanagama Hoser, 2015), or: 2019. The Linz Zoocode project: a set of new proposals 3/ Tympanum hidden; covered by skin; body scales are strongly regarding the terminology, the Principles and Rules of zoological heterogeneous, many of the larger scales on the body and head nomenclature. First report of activities (2014-2019). Bionomina keeled or moderately spinose; no series of enlarged, spinose (online), 17:1-111. scales on either side of the base of the tail (subgenus Even, E. 2005. Reptielen zoeken in Australië. Lacerta 63(2):48- Turnbullagama Hoser, 2015). 65. Notactenophorus Hoser, 2015 is readily separated from all other Fitzinger, L . J. 1843. Systema reptilium. Vindobonae [Wien] members of the genus Ctenophorus (where it has been placed :Braumüller et Seidel,1843. until now, as defined in Cogger 2014) and Pseudoctenophorus Glauert, L. 1959. Herpetological Miscellanea. XI. Dragon Lizards Hoser, 2015 by the following unique suite of characters: of the Genus Amphibolurus. West. Aust. Naturalist 7(2):42-51. Tympanum is hidden being covered by skin, the body scales are smooth, mostly small, homogenous, with scattered larger but Gray, J. E. 1845. Catalogue of the specimens of lizards in the small, flat scales, not keeled or spinose, with a dorsal pattern of a collection of the British Museum. Trustees of the British Museum/ longitudinal dorso-lateral series of five or six large black spots on Edward Newman, London:UK:xxvii+289 pp. either side. Günther, A. 1875. A list of the saurians of Australia and New Distribution: C. shireenhoserae sp. nov. is known only from Zealand. pp. 9-19. In: Richardson, J. and Gray, J. E. The zoology near the type locality and nearby areas in Western Australia, of the voyage of H.M.S. Erebus and Terror, during the years generally between Paynes Find and Mount Magnet in Western 1839 to 1843. By authority of the Lords Commissioners of the Australia. Admiralty. Vol. 2. E. W. Janson, London, UK. Hoser, R. T. 1989. Australian Reptiles and Frogs. Pierson and C. sharonhoserae sp. nov. is found in the semi-arid zone of Co., Mosman, NSW, 2088:238 pp. south-western Western Australia, away from the Darling Range and south coast of Western Australia, and also south of the Hoser, R. T. 1991. Endangered Animals of Australia. Pierson mulga-eucalypt line in south-west Australia. The distribution is Publishing, Mosman, NSW, Australia:240 pp. mainly in the Goldfields region of southern Western Australia, Hoser, R. T. 1993. Smuggled: The Underground Trade in Australia. Australia’s Wildlife. Apollo Publishing, Moss Vale, NSW, C ornatus is restricted to the Darling Range (near Perth) and Australia:149 pp. wetter parts of the south coast of south-western Australia. Hoser, R. T. 1996. Smuggled-2: Wildlife Trafficking, Crime and Etymology: C. shireenhoserae sp. nov. is named in honour of Corruption in Australia. Kotabi Publishing, Doncaster, Victoria, my wife Shireen Vanessa Hoser from a remote part of Africa Australia:280 pp. called “Athlone”, in recognition of her contributions to herpetology Hoser, R. T. 2007. Wells and Wellington - It’s time to bury the over more than 20 years. hatchet. Calodema Supplementary Paper 1:1-9.

Hoser, R. T. 2009. Creationism and contrived science: A review of Kaiser, H. 2014a. Comments on Spracklandus Hoser, 2009 recent python systematics papers and the resolution of issues of (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the taxonomy and nomenclature. Australasian Journal of availability of the generic name and for the nomenclatural Herpetology 2:1-34. (3 February). validation of the journal in which it was published. Bulletin of Hoser, R. T. 2012a. Exposing a fraud! Afronaja Wallach, Wüster Zoological Nomenclature, 71(1):30-35. and Broadley 2009, is a junior synonym of Spracklandus Hoser Kaiser H. 2014b. Best Practices in Herpetological Taxonomy: 2009! Australasian Journal of Herpetology 9 (3 April 2012):1-64. Errata and Addenda. Herpetological Review, 45(2):257-268. Hoser, R. T. 2012b. Robust taxonomy and nomenclature based Kaiser, H., Crother, B. L., Kelly, C. M. R., Luiselli, L., O’Shea, M., on good science escapes harsh fact-based criticism, but remains Ota, H., Passos, P., Schleip, W. D. and Wüster, W. 2013. Best unable to escape an attack of lies and deception. Australasian practices: In the 21st Century, Taxonomic Decisions in Journal of Herpetology 14:37-64. Herpetology are Acceptable Only When supported by a body of Hoser, R. T. 2013. The science of herpetology is built on Evidence and Published via Peer-Review. Herpetological Review evidence, ethics, quality publications and strict compliance with 44(1):8-23. the rules of nomenclature. Australasian J. of Herp. 18:2-79. Kinghorn, J. R. 1924. Reptiles and batrachians from south and Hoser, R. T. 2015a. Dealing with the “truth haters” ... a summary! south-west Australia. Records of the Australian Museum Introduction to Issues 25 and 26 of Australasian Journal of 14(3):163-183. Herpetology. Including “A timeline of relevant key publishing and Lebas, N. R. and Spencer, P. B. S. 2000. Polymorphic other events relevant to Wolfgang Wüster and his gang of microsatellite markers in the Ornate Dragon lizard, Ctenophorus thieves.” and a “Synonyms list”. Australasian Journal of ornatus. Molecular Ecology 9(3):365-366. Herpetology 25:3-13. Melville, J., Schulte, J. A. and Larson, A. 2001. A molecular Hoser, R. T. 2015b. The Wüster gang and their proposed “Taxon phylogenetic study of ecological diversification in the Australian Filter”: How they are knowingly publishing false information, lizard genus Ctenophorus. Journal of Experimental Zoology: recklessly engaging in taxonomic vandalism and directly Molecular and Developmental Evolution 291(4):339-353. attacking the rules and stability of zoological nomenclature. Melville, J., Haines, M. L., Hale, J., Chapple, S. and Ritchie, E. G. Australasian Journal of Herpetology 25:14-38. 2016. Concordance in phylogeography and ecological niche Hoser, R. T. 2015c. Best Practices in herpetology: Hinrich modelling identify dispersal corridors for reptiles in arid Australia. Kaiser’s claims are unsubstantiated. Journal of Biogeography, 10 June (online). Australasian Journal of Herpetology 25:39-64. Peters, W. C. H. 1876. Über die von S. M. S. Gazelle Hoser, R. T. 2015d. PRINO (Peer reviewed in name only) mitgebrachten Amphibien. Monatsber. königl. Akad. Wiss. Berlin. journals: When quality control in scientific publications fails. 1876 (August):528-535. Australasian Journal of Herpetology 26:3-64. Pyron, R. A., Burbrink, F. T. and Weins, J. J. 2013. A phylogeny Hoser, R. T. 2015e. Rhodin et al. 2015, Yet more lies, and revised classification of Squamata, including 4161 species of misrepresentations and falsehoods by a band of thieves intent on lizards and snakes. Published online at: stealing credit for the scientific works of others. Australasian http://www.biomedcentral.com/1471-2148/13/93. Journal of Herpetology 27:3-36. Ride, W. D. L. (ed.) et al. (on behalf of the International Hoser, R. T, 2015f. Comments on Spracklandus Hoser, 2009 Commission on Zoological Nomenclature) 1999. International (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the code of Zoological Nomenclature. The Natural History Museum - availability of the generic name and for the nomenclatural Cromwell Road, London SW7 5BD, UK (also commonly cited as validation of the journal in which it was published (Case 3601; “The Rules”, “Zoological Rules” or “ICZN 1999”). see BZN 70: 234-237; comments BZN 71:30-38, 133-135). Sadlier, R. A., Colgan, D., Beatson, C. A. and Cogger, H. G. Australasian Journal of Herpetology 27:37-54. 2019. Ctenophorus spinodomus sp. nov., a New Species of Hoser, R. T. 2015g. Australian agamids: Eighteen new species Dragon Lizard (Squamata: Agamidae) from Triodia Mallee from the genera Amphibolurus Wagler, 1830, Lophognathus Habitat of Southeast Australia. Rec. Aust. Mus. 71(5):199-215. Gray, 1842, Rankinia Wells and Wellington, 1984, Diporiphora Sternfeld, R. 1925. Beiträge zur Herpetologie Inner-Australiens. Gray, 1842, Tympanocryptis Peters, 1863, as well as three new Abhandlungen Herausgegeben von der Senckenbergischen genera and six new subgenera. Australasian J. Herp. 30:37-64. Naturforschenden Gesellschaft, 38:221-251. Hoser, R. T. 2019a. 11 new species, 4 new subspecies and a Storr, G. M. 1967. Geographic races of the agamid lizard subgenus of Australian Dragon Lizard in the genus Amphibolurus caudicinctus. J. Royal Soc. of WA. 50:49-56. Published in hard copy on 25 April 2020 Tympanocryptis Peters, 1863, with a warning on the conservation Storr, G. M. 1981. Three new agamid lizards from Western status and long-term survival prospects of some newly named Australia. Rec. of the Western Australian Museum 8(4):599-607. taxa. Australasian Journal of Herpetology 39:23-52. Storr, G. M., Smith, L. A. and Johnstone, R. E. 1983. Lizards of Hoser, R. T. 2019b. Richard Shine et al. (1987), Hinrich Kaiser et Western Australia 2: dragons and Monitors. Western Australian al. (2013), Jane Melville et al. (2018 and 2019): Australian Museum, Perth, Western Australia, Australia:114 pp. Agamids and how rule breakers, liars, thieves, taxonomic vandals Victorian Civil and Administrative Tribunal (VCAT). 2015. Hoser v and law breaking copyright infringers are causing reptile species Department of Environment Land Water and Planning (Review to become extinct. Australasian Journal of Herpetology 39:53-63. and Regulation) [2015] VCAT 1147 (30 July), judgment and Hoser, R. T. 2020. A long overdue refinement of the taxonomy of transcript). the Mallee Dragon Complex Ctenophorus (Phthanodon) fordi Wells, R. W. and Wellington, C. R. 1985. A classification of the (Storr, 1965) sensu lato with the formal descriptions of four new Amphibia and Reptilia of Australia. Australian Journal of subspecies. Australasian Journal of Herpetology 43:41-49. Herpetology Supplementary Series 1:1-61. Kaiser, H. 2012a. SPAM email sent out to numerous recipients on Wilson, S. K. and Knowles, D. G. 1988. Australia’s reptiles. A 5 June 2012. photographic reference to the terrestrial reptiles of Australia. Kaiser, H. 2012b. Point of view. Hate article sent as attachment Collins Australia, Sydney, Australia:447 pp. with SPAM email sent out on 5 June 2012. Wilson, S. and Swan, G. 2017. A complete guide to reptiles of Kaiser, H. 2013. The Taxon Filter, a novel mechanism designed Australia, 3rd ed. Chatswood: New Holland, 647 pp. to facilitate the relationship between taxonomy and nomenclature, CONFLICTS OF INTEREST vis-à-vis the utility of the Code’s Article 81 (the Commission’s plenary power). Bulletin of Zoological Nomenclature 70(4) None. December 2013:293-302. Hoser 2020 - Australasian Journal of Herpetology 42:50-64. Available online at www.herp.net Copyright- Kotabi Publishing - All rights reserved